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PubMed2006

Heterosis analysis of Haemonchus contortus resistance and production traits in Rhoen sheep, Merino Land sheep and crossbred lambs.

A crossbreeding program was conducted to evaluate the resistance status to Haemonchus contortus in reciprocal crossbred (F(1)) lambs and their Merino Land sheep (Ml) and Rhoen sheep (Rh) purebred counterparts. A total of 406 lambs were included in the study. Faecal egg counts (FEC) and haematocrit values (Hc) of all the lambs were collected 4 and 8 weeks after an artificial infection with H. contortus. FAMACHA scores were given at the same time. Worm counts of all lambs were obtained after slaughtering at 21 weeks of age. The Ml group consistently showed the highest body weights, daily weight gains and best carcass performances, whereas the Rh group showed the lowest results. Most of the body weight parameters in the F(1) group fell intermediately between the Ml and Rh groups, though the heterosis analysis of body weight and carcass parameters showed a tendency to favour the crossbreeding group Rh x Ml. The Rh group showed, on average, the highest FEC and the lowest Hc values compared to the Ml. However, the nematode counts were not significantly different between the pure breeds. In comparison, the worm counts of the Rh x Ml crosses were significantly different from those in the Ml x Rh crosses. A heterosis analysis showed that FEC, Hc and the worm count in the F(1) group favoured the crossbreeding group Rh x Ml. These results suggested that crossbreeding Rh to Ml sheep may be a suitable way of producing lambs with improved resistance to H. contortus infection without any negative effects on production traits.

Animals↗

PubMed2006

Effect of sericea lespedeza (Lespedeza cuneata) fed as hay, on natural and experimental Haemonchus contortus infections in lambs.

Parasitic infection is one of the leading economic constraints in small ruminant production. This problem is exacerbated as the resistance of nematode populations to chemical treatment (anthelmintics) becomes increasingly more common. Condensed tannin (CT) containing plants are being investigated as alternative solutions to address these problems. This investigation was conducted to evaluate the effect of the CT containing forage, Lespedeza cuneata (sericea lespedeza, SL) fed as hay, on Haemonchus contortus infection in sheep. Naturally and experimentally infected lambs were fed either SL or bermudagrass (BG, control) hay for 49 days. All lambs were fed BG hay for an additional 14 days. SL hay effectively reduced (67-98%) fecal egg count (FEC) during the time of feeding for both infection groups. FEC increased in both infection groups after SL feeding was stopped which indicated an effect on fecundity. SL hay feeding also reduced worm numbers, with more of an effect on reducing naturally infected worm burdens (67.2%) than on establishment of incoming larvae (26.1%). SL fed as hay may be more useful to remove existing worms than establishing worms. The decrease in FEC would have the benefit of reduced pasture contamination.

Animal Feed↗

PubMed2006

Effect of sainfoin (Onobrychis viciifolia) silage and hay on established populations of Haemonchus contortus and Cooperia curticei in lambs.

The objective of the study was to examine the effect of dried and ensiled sainfoin (Onobrychis viciifolia) on established populations of Haemonchus contortus (abomasum) and Cooperia curticei (small intestine) in lambs under controlled conditions. Twenty-four parasite naïve lambs were inoculated with a single dose of infective larvae of these parasites 28 days prior to the start of the feeding experiment. Twenty-four days post-infection, 4 days prior to the start of the feeding experiment, animals were allocated to four groups according to egg excretion, live weight and sex. Groups A and B received sainfoin hay and control hay, respectively, for 16 days. Groups C and D were fed on sainfoin silage or control silage for the same period. Feeds were offered ad libitum and on the basis of daily refusals were supplemented with concentrate in order to make them isoproteic and isoenergetic. Individual faecal egg counts on a dry matter basis (FECDM) were performed every 3-4 days and faecal cultures and packed cell volume (PCV) measurements were done weekly. After 16 days of experimental feeding, all animals were slaughtered and adult worm populations were determined. The consumption of conserved sainfoin was associated with a reduction of adult H. contortus (47% in the case of hay, P<0.05; 49% in the case of silage, P=0.075) but had little effect on adult C. curticei. Compared to the controls, H. contortus specific FECDM was reduced by 58% (P<0.01) in the sainfoin hay group and by 48% (P=0.075) in the sainfoin silage group. For both sainfoin feeds FECDM specific to C. curticei were significantly decreased when compared to the control feeds (hay 81% and silage 74%, both tests P<0.001). Our data suggest that different mechanisms were responsible for the reduction in FECDM in response to feeding tanniferous fodder. For H. contortus, the decrease seemed to be due to a nematocidal effect towards adult H. contortus. In contrast for C. curticei, the reduction in FECDM appeared to be a result of a reduced per capita fecundity. For both, hay and silage, an antiparasitic effect could be shown, offering promising perspectives for the use of conserved tanniferous fodder as a complementary control approach against GIN.

Abomasum↗

PubMed2006

Phenotyping and genotyping of Haemonchus contortus isolates reveals a new putative candidate mutation for benzimidazole resistance in nematodes.

In order to monitor and eventually control the spread of drug-resistant Haemonchus contortus, knowledge of the molecular mechanisms underlying resistance is essential. Here we phenotypically and genotypically characterize three multidrug-resistant H. contortus field isolates from Australia and South Africa. All were significantly less susceptible to thiabendazole than a sensitive reference strain in an in vitro egg-hatch assay. The sensitivity was further reduced in a surviving population after treatment of infected sheep with albendazole. The beta-tubulin genes were amplified from genomic DNA of the H. contortus isolates, cloned, and sequenced. There was a high degree of sequence variation. The known mutation phenylalanine-200 to tyrosine (F200Y) occurred in 60% of the sequences from resistant isolates, but not in the sensitive reference. Interestingly, 90% of the beta-tubulin sequences from resistant isolates lacking tyrosine-200 carried another mutation nearby, glutamate-198 to alanine (E198A). This mutation was not found in the sensitive isolate, nor in sequences from resistant isolates carrying the mutation F200Y. However, the mutation E198A is known from benomyl-resistant isolates of phytopathogenic fungi such as Monilinia fructicola. The finding that alanine-198 correlates with thiabendazole resistance in H. contortus isolates from South Africa and Australia suggests that also in nematodes, the mutation E198A plays a role in benzimidazole resistance. Alanine-198 alleles of beta-tubulin can be detected by PCR-RFLP and we suggest to include this test in future surveys of H. contortus field populations.

Amino Acid Sequence↗

PubMed2006

Detection and measurement of benzimidazole resistance alleles in Haemonchus contortus using real-time PCR with locked nucleic acid Taqman probes.

Benzimidazole resistance is a common problem in parasitic nematodes of ruminants and early detection is vital if its spread is to be monitored and controlled. Real time PCR offers a fast and reliable method for rapid detection and measurement of resistance allele frequencies. In Haemonchus contortus a single nucleotide polymorphism at codon 200 of the beta-tubulin gene (TTC to TAC), causing a phenylalanine to tyrosine amino acid substitution, has been shown to be involved in many cases of resistance. Locked nucleic acid (LNA) Taqman probes have been used in this work to detect and measure the frequency of resistance alleles in individual and multiple H. contortus. Detection of resistant genotypes using LNA Taqman probes in individual H. contortus is simpler and more reliable than with previously described assays. Measurement of the frequency of resistant alleles in populations of H. contortus was achieved by using the cycle threshold (C(t)) values and a standard curve derived from populations with known allele frequencies. Results using the LNA probes on individual and multiple worms gave similar results to the allele specific PCR. The sensitivity of the LNA assay on multiple nematodes allowed reliable detection of > or = 10% resistance allele frequency. Using the final fluorescence method, it was possible to differentiate populations with approximately 0, 5 and 10% resistance allele frequencies.

Alleles↗

PubMed2006

Recombinant galectins of male and female Haemonchus contortus do not hemagglutinate erythrocytes of their natural host.

Recombinant galectins of female and male adult worms of Haemonchus contortus were expressed in Escherichia coli and their hemagglutinating activities to human and different animal erythrocytes were analyzed. The results showed that female and male galectins could be highly expressed in E. coli using a temperature-sensitive plasmid, with the recombinant protein being mainly appeared in inclusion bodies. Hemagglutinating activity assays showed that both of the galectins hemagglutinated human A, B, O type, dog, rabbit, chicken and mouse erythrocytes at the high concentration of 40 microg/well, but did not hemagglutinate erythrocytes of the natural host of H. contortus, the goat. Sugar inhibition assays confirmed that, out of eight sugars tested, only lactose was effective to inhibit agglutination of human type B erythrocytes by the recombinant galectins.

Animals↗

PubMed1997

Availability of Haemonchus contortus L3 larvae on pasture at Kiboko: a semi-arid warm agro-climatic zone in Kenya.

A study was carried out to determine the availability of Haemonchus contortus L3 larvae on pasture in a semi-arid warm agro-climatic zone of Kenya. By means of tracer sheep, it was shown that no H. contortus L3 larvae were available on pasture during the dry periods of the year (July-October and February). They were only available on pasture during the rainy season (November-January and March-June). Sheep permanently grazed on the same pasture however, harboured adult H. contortus in their abomasa throughout the year indicating that the perpetuation of haemonchosis in livestock in this agro-climatic zone was greatly dependent on the ability of the parasite to survive in the host throughout all seasons.

Animals↗

PubMed2002

A glutamate-gated chloride channel subunit from Haemonchus contortus: expression in a mammalian cell line, ligand binding, and modulation of anthelmintic binding by glutamate.

Glutamate-gated chloride channels (GluCls) are inhibitory ion channels that are sensitive to the antiparasitic drugs ivermectin (IVM) and moxidectin (MOX). We have transiently transfected COS-7 cells with a subunit of a GluCl (HcGluCla) from the parasitic nematode Haemonchus contortus. This subunit bound [3H]-IVM and [3H]-MOX with K(d) values of 0.11+/-0.021 and 0.18+/-0.02nM, respectively. Displacement analysis revealed that IVM and MOX bind to the same site on HcGluCla and that this site is likely distinct from the glutamate binding site. Glutamate was found to be an allosteric modulator of [3H]-MOX and [3H]-IVM binding and increased the affinity of [3H]-MOX for HcGluCla by more than 50% and that of [3H]-IVM by more than 7-fold. These results point to both similarities and differences in the interactions of IVM and MOX with the GluCl. Aspartate, which is structurally similar to glutamate, had little or no effect on [3H]-IVM and [3H]-MOX binding, suggesting that this ligand does not induce the conformational change necessary to potentiate macrocyclic lactone binding. These results also indicate that it may be possible to enhance the efficacy of macrocyclic lactone anthelmintics by administering these compounds with ligands acting allosterically to enhance their binding.

Animals↗

PubMedDate not supplied

The immunogenetics of resistance to Trichostrongylus colubriformis and Haemonchus contortus parasites in sheep.

Three possible immunogenetic markers for resistance to intestinal parasites in sheep have been studied. Allotypes of the major histocompatibility complex (MHC) of the sheep have been investigated as markers, using serological typing or gene probes, for associations between allotypes and resistance to parasites in selected high responder and low responder lines of sheep. Only the serologically-determined class I ovine leucocyte antigen (OLA) types SY 1a and SY 1b have been found to be consistently associated with increased resistance to Trichostrongylus colubriformis, but this association has not extended to the immunologically distinct Haemonchus contortus parasite. Gene probes of the sheep DRB, DQB and DQA MHC class II loci have detected animals with increased susceptibility to T. colubriformis. Eosinophilia was investigated as a marker and found to be associated with increased resistance to parasites in lines of Australian Merinos and New Zealand Romneys selected for resistance on the basis of low faecal egg count. Blood eosinophilia was distinct from eosinophil infiltration of the gut which was poorly associated with resistance. The mechanism of parasite resistance appeared to involve the release of vasoactive amines and leukotrienes into intestinal mucus, since the selected high responder sheep to T. colubriformis and H. contortus had significantly increased amounts of these agents in their gut mucus, compared with selected low responder or random-bred sheep. Antibodies to T. colubriformis and H. contortus have also been used as markers to select high responder sire groups of lambs in contact with the parasites, for the first time, at weaning. This assay had the advantage of detecting distinct antigens for the two parasites, which would allow resistance to the species of parasite to be selected in the lambs. Vaccines have been developed against H. contortus using 'novel' gut antigens from the parasite, but variable responsiveness of the host sheep seemed to result in varying degrees of protection which were stimulated by these vaccines.

Animals↗

PubMed2002

Distinct characteristics of two intestinal protein compartments discriminated by using fenbendazole and a benzimidazole resistant isolate of Haemonchus contortus.

The intestine of Haemonchus contortus is hypersensitive to the effects of the anthelmintic fenbendazole (FBZ). The effects are postulated to stem from disruption of microtubules and interference with apical secretory vesicle transport, followed by release of digestive enzymes into the intestinal cell cytoplasm. Here, FBZ caused marker proteins for both apical (pepsinogen-like protease, PEP-1) and basal (cystatin-like protein) protein compartments to became homogeneously distributed in the cytoplasm of H. contortus intestinal cells. The observations with PEP-1 support the hypothesis that release of hydrolytic enzymes into the intestinal cell cytoplasm contributes to the mechanism of benzimidazole efficacy. A benzimidazole resistant isolate of H. contortus expressed type 1 and 2 intestinal beta-tubulin transcripts that would encode predominantly tyr200 and phe200 variants, respectively. This isolate was resistant to the known intestinal cell alterations induced by FBZ treatment in the susceptible isolate, including inhibition of apical vesicle transport. These results implicate type 1 beta-tubulin in mediating apical vesicle transport in intestinal cells and suggest that the tyr200 variant is a determinant of FBZ resistance in intestinal cells. In contrast, the basal protein compartment demonstrated sensitivity to FBZ treatment in these otherwise "resistant" worms. Hence, distinct FBZ-sensitive components appear to be involved in distributing intestinal proteins into the described apical and basal compartments of normal worms.

Animals↗

PubMedDate not supplied

Non-classic characteristics define prominent DNase activities from the intestine and other tissues of Haemonchus contortus.

The anthelmintic fenbendazole (FBZ) induces nuclear DNA fragmentation (DF) in intestinal cells of Haemonchus contortus. The DNA fragments had 3'-OH, which suggests involvement of a neutral DNase. To identify candidate DNase(s) involved, DNase activity in H. contortus intestine and other worm fractions was characterized relative to classic DNases I (neutral) and II (acidic). Seven distinct DNase activities were identified and had Mrs of 34, 36, 37 or 38.5 kDa on zymographic analysis. The different activities were distinguished according to pH requirement, sensitivity to 10 mM EDTA and worm compartment. Activities of intestinal DNases at 34, 36 and 38.5 kDa were sensitive to EDTA at pH 5.0 and 7.0. Sensitivity to EDTA at pH 5.0 was unexpected compared to classic acidic DNase II activity, suggesting unusual properties of these DNases. In whole worms, however, the activities at 36 and 38.5 kDa were relatively insensitive to EDTA, indicating predominance of DNases that are distinct from the intestine. The activity at 37 kDa in excretory/secretory products had an acidic pH requirement and was insensitive to EDTA, resembling classic acidic DNase activity. Under conditions of pH 5.0 and 7.0, intestinal DNases produced 3'-ends that could be labeled by terminal deoxynucleotidyl transferase, indicating presence of 3'-OH. The labeling of 3'-ends at pH 5.0, again, was unexpected for acidic DNase activity. These results and several other activities suggest that multiple H. contortus DNases have characteristics distinct from the classic mammalian DNases I and II. Treatment of H. contortus with FBZ did not induce any detectable DNase activities distinct from normal intestine, although relative activities of intestinal DNases appear to have been altered by this treatment.

Animals↗

PubMed1999

Core alpha1-->3-fucose is a common modification of N-glycans in parasitic helminths and constitutes an important epitope for IgE from Haemonchus contortus infected sheep.

Synthesis of parasite specific IgE plays a critical role in the defence against helminth infections. We report here that IgE from serum from Schistosoma mansoni infected mice and Haemonchus contortus infected sheep recognizes complex-type N-glycans from Arabidopsis thaliana, which contain R-GlcNAcbeta1-->4(Fucalpha1-->3)GlcNAcbeta1-Asn (core alpha1-->3-Fuc) and Xylbeta1-->2Manbeta1-->4GlcNAcbeta1-R (core beta1-->2-Xyl) modifications, and honeybee phospholipase A2, which carries N-glycans that contain the core alpha1-->3-Fuc epitope. Evidence is presented that core alpha1-->3-fucosylated N-glycans bind a substantial part of the parasite specific IgE in serum of H. contortus infected sheep. These results suggest that the core alpha1-->3-Fuc antigen may contribute to induction of a Th2 response leading to the production of IgE. In addition we show here that N-glycans carrying core alpha1-->3-Fuc and beta1-->2-Xyl antigens are synthesized by many parasitic helminths and also by the free living nematode Caenorhabditis elegans. Since N-glycans containing the core alpha1-->3-Fuc have also been implicated in honeybee and plant induced allergies, this conserved glycan might represent an important common IgE epitope.

Animals↗

PubMed2001

Characterisation of Haemonchus contortus-derived cell populations propagated in vitro in a tissue culture environment and their potential to induce protective immunity in sheep.

Cell populations derived from viable Haemonchus contortus L(3) larvae were propagated in vitro in a tissue culture environment for a prolonged period (>48 months). Microscopic evaluation of H. contortus-derived cell populations revealed gross morphological characteristics highly analogous to those described for cell types originating from species of plant nematodes propagated in vitro in a tissue culture environment for a briefer period of time (<6 months). The characterisation of extracts harvested from tissue culture populations of H. contortus-derived cells by SDS-PAGE analysis detected molecular fractions of approximately 29, 45, 55, and 200-kDa that closely correlated with reports for preparations obtained from intact/viable H. contortus larvae. Complementary investigations detected the dual biochemical expression of phosphohydrolase and aminopeptidase-M activities based on the hydrolysis of the synthetic enzyme-specific substrates, para-nitrophenylphosphate and leucine-para-nitroanaline, respectively. The identification of phosphohydrolase and aminopeptidase-M-like biochemical activity in fractions harvested from H. contortus-derived cell populations and propagated in vitro in tissue culture served as evidence validating their parasitic-origin. Further validation of H. contortus-derived cell populations propagated in tissue culture entailed the formulation of Triton X-100 extracts containing potential immunoprotective antigens with SEAM adjuvant and its administration by intramuscular injection (100 microg total protein) to healthy sheep (n=8) on day 0 (left rear-limb) and day +14 (right rear-limb). Animals on day 28 subsequently received a single oral challenge of 10,000 infective L(3)-stage H. contortus larvae. Applying ELISA methodologies, increases in antigen-specific IgM and IgG were detected in ovine serum samples. Interpretation of experimental findings revealed that sheep with the greatest antigen-specific humoral immune responses (IgG titre 1/3125) also demonstrated a degree of reduced abomasal H. contortuslarvae burdens (60% reduction). Polyclonal antibody from immunoprotected sheep was subsequently found to recognise both the: (i), digestive tract; and (ii), antigen extracts associated with intact/viable H. contortus larvae. These experimental findings reveal the potential feasibility of propagating parasite-derived cell populations in an in vitro tissue culture environment in a manner that retains their ability to express immunoprotective antigenic fractions.

Abomasum↗

PubMed2001

Amplified fragment length polymorphism analysis of genetic diversity of Haemonchus contortus during selection for drug resistance.

For the first time we used amplified fragment length polymorphism on individual nematode parasites to analyse the genetic diversity between and within isolates during consecutive stages of increased benzimidazole resistance and of increased levamisole resistance of Haemonchus contortus. The genetic diversity of the H. contortus genome turned out to be unusually high, within and between the isolates. The difference between individuals of an isolate could be as high as between individuals of two different mammalian species that do not interbreed. During benzimidazole selection the genetic constitution of the population was changed, but did not lead to a decrease in the genetic diversity. The selection for levamisole resistance resulted in a limited reduction of the genetic diversity only after the first selection step. The extensive genetic diversity apparently has allowed a fast and flexible response of H. contortus to drug selection as shown by the appearance of drug resistant isolates. This selection however has little or no effect on the extent of the genetic diversity of these resistant isolates. Implications for more sustainable control methods are discussed.

Animals↗

PubMed2001

Analysis of developmentally regulated genes of the parasite Haemonchus contortus.

Differential regulation of gene expression in the development of Haemonchus contortus was analysed using RNA arbitrarily-primed PCR. A study of third-stage larval and adult H. contortus revealed large differences between the two stages; 32 and 30% unique third-stage larval and adult RNA arbitrarily-primed PCR products, respectively. This finding is consistent with a high degree of differential gene expression between these developmental stages. A number of adult products were sequenced, revealing 11 molecules to be similar to deposits within sequence databases. Four other molecules that did not have significant similarity to sequences in the databases may represent developmentally regulated genes specific to H. contortus. Northern analysis of the putative adult-expressed molecules with homologues in the databases confirmed that four were expressed only in adults, while four were expressed in both stages, but had different sized transcripts. This may reflect differential splicing, or expression of closely related but different molecules at different life cycle stages. Two molecules were present in mRNA populations from both stages, suggesting these were false stage-associated molecules. No transcript was detected for one molecule by Northern analysis, probably due to low level of expression. In situ hybridisation analysis was used to localise expression of transcripts in the adult parasite, in particular, to gain some insight into the nature of those molecules with no known predicted function.

Amino Acid Sequence↗

PubMed2002

HcSTK, a Caenorhabditis elegans PAR-1 homologue from the parasitic nematode, Haemonchus contortus.

A putative serine/threonine protein kinase (HcSTK) from the parasitic nematode Haemonchus contortus was characterised at the mRNA and amino acid levels. HcSTK displays a high level of identity (85-93% in the catalytic domain) with proteins of the PAR-1/MARK serine/threonine protein kinase (STK) subfamily, which represent signal transduction molecules involved in establishing and maintaining polarity in proliferating and differentiating cells. The transcript of hcstk is expressed in different developmental stages (second-, third-, fourth-stage larvae and adults) and various organs (muscle, intestine and reproductive) of H. contortus. In addition, there are several isoforms which appear to relate to a single gene. The expression profile of hcstk is similar to that of Caenorhabditis elegans PAR-1, and the level of sequence identity among members of the PAR-1/MARK STK subfamily, representing a range of species of vertebrates (e.g. humans and rodents), invertebrates (e.g. insects and C. elegans) and yeast, suggests that HcSTK may be involved in a conserved signal transduction pathway.

Amino Acid Sequence↗

PubMed2003

Catalase induction protects Haemonchus contortus against hydrogen peroxide in vitro.

The response of Haemonchus contortus to oxidative stress in vitro was examined by measuring catalase activities in adult and L4 stage worms exposed to hydrogen peroxide generated by a glucose/glucose oxidase system. Adult nematodes showed increases of up to 2.3-fold in catalase activity after 42 h exposure to the peroxide. L4 nematodes showed up to 4.6-fold induction. A two-stage dose-response was apparent, with catalase activities increasing as the peroxide levels increased, before a return to control levels at higher peroxide concentrations, most likely reflecting a balance between induction and toxicity of the inducing agent itself. Adult nematodes exposed to low levels of peroxide for 24h (hence, having enhanced catalase activities) showed an ability to tolerate subsequent exposure to toxic levels of the peroxide compared to worms with no pre-exposure. An increase of up to approximately threefold in the LC(50) of the hydrogen peroxide generating system was observed after hydrogen peroxide pre-exposure. This indicates that exposure to low oxidant levels lead to an increase in defensive enzyme activities, which allows the nematode to survive subsequent oxidant threats more effectively. The ability of H contortus to increase its catalase activity may be crucial in allowing it to respond to the production of reactive oxygen species by host phagocytes in vivo.

Animals↗

PubMed2003

The nature and prospects for gut membrane proteins as vaccine candidates for Haemonchus contortus and other ruminant trichostrongyloids.

Substantial progress has been made in the last decade in identifying several antigens from Haemonchus contortus which, in their native form, stimulate useful levels of protective immunity (70-95% reductions in faecal egg output) in the ovine host. Much work has focussed on proteins/protein complexes expressed on the surface of the worm gut which are exposed to the blood meal, and, hence, antibody ingested with it. The antigens generally, but not in all cases, show protease activity and antibody is thought to mediate protective immunity by blocking the activity of enzymes involved in digestion within the worm. This review summarises the protective efficacy, as well as the biochemical and molecular properties, of the principal candidate antigens which are expressed in the gut of these parasites. Of course, such antigens will have to be expressed as recombinant proteins to be sufficiently cost-effective for use in a commercial vaccine and the current status of recombinant antigen expression is discussed with particular reference to conformation and glycosylation. There is a need for continued antigen definition even in the confines of gut antigens and potential targets can be selected from the rapidly expanding genome/EST datasets on the basis of predicted functional homology. Gene knockout technologies such as RNA interference have the potential to provide high throughput, rapid and inexpensive methods to define whether the protein product of a particular gene would be a suitable vaccine candidate.

Animals↗