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We tested the hypothesis that Haemonchus contortus and Trichostrongylus colubriformis would adapt to long-term exposure to sheep that were either genetically resistant or susceptible to H. contortus. Sheep genotypes were from lines with 10 years prior selection for low (resistant, R) or high (susceptible, S) faecal worm egg count (WEC) following H. contortus infection. Long-term exposure of H. contortus and T.colubriformis to R or S genotypes was achieved using serial passage for up to 30 nematode generations. Thus, we generated four nematode strains; one strain of each species solely exposed to R sheep and one strain of each species solely exposed to S sheep. Considerable host genotype differences in mean WEC during serial passage confirmed adequate nematode selection pressure for both H. contortus (R 4900 eggs per gram (epg), S 19,900 epg) and T. colubriformis (R 5300 epg, S 13,500 epg). Adaptation of nematode strain to host genotype was tested using seven cross-classified tests for H. contortus, and two cross-classified and one outbred genotype test for T. colubriformis. In the cross-classified design, where each strain infects groups of R, S or randomly bred control sheep, parasite adaptation would be indicated by a significant host genotype by nematode strain interaction for traits indicating parasite reproductive success; specifically WEC and, for H. contortus strains, packed cell volume. We found no significant evidence of parasite adaptation to host genotype (P>0.05) for either the H. contortus or T. colubriformis strains. Therefore, we argue that nematodes will not adapt quickly to sheep bred for nematode resistance, where selection is based on low WEC, although selecting sheep using a subset of immune functions may increase adaptation risk. Our results support the hypothesis that nematode resistance is determined by many genes each with relatively small effect. In conclusion, selection of sheep for nematode resistance using WEC should be sustainable in the medium to long-term.

Breeding for disease resistance in Merino sheep in Australia has attracted considerable research and development attention. Increased labour costs, the reduced efficacy of common anthelmintics and insecticides, consumer demand for products which are free of chemicals and the poor prognosis of alternative long-term control strategies are all forcing sheep breeders to contemplate the best animal health options available, including selective breeding for resistance. The three major diseases which affect sheep production include gastrointestinal nematode parasites, flystrike (cutaneous myiasis) and footrot. Genetic improvement has been shown to be feasible in all these diseases, possibly with little adverse effect on genetic progress in other production traits. Programmes for resistance to internal parasites to be included in practical breeding programmes are now in progress. This paper deals with the incentives for focusing on the three major diseases in Merino sheep, the potential sources of genetic variation, and the means to exploit these sources of variation. The authors also highlight gains and benefits achieved in experimental selection flocks, and the difficulties and options available for commercial breeders.

When experiments are established to compare parasitological differences between genetic groups of animals, sampling of families within groups and individuals within families are both important. Many parasitological measurements are quite variable, so relatively large numbers of animals are required to detect small differences. Calculations are presented which show, for example, that 100 animals per genetic group will only have an 80% chance of detecting a difference in faecal worm egg counts as significant (P < 0.05) if the real difference between the groups is about 3-fold. When sampling genetic groups such as flocks or breeds, it is preferable to represent as many families as possible.

Resistance to nematode parasites can be improved by selection, but efforts to include appropriate traits in commercial livestock breeding programs are only a recent development. Procedures for including resistance in breeding programs are similar to those involving other traits. The steps are described with special reference to sheep, and areas are highlighted where particular considerations exist. Three approaches are described and contrasted: breeding for resistance (reduced parasite numbers, as determined by faecal worm egg count); resilience (production during parasitism); or number of treatments required during parasitism. It is necessary, but difficult, to assess the economic benefits of improving resistance relative to other traits. Disease costs vary widely depending on the prevalence of the disease and on the availability, effectiveness and sustainability of alternative control measures. Costs of treatment and control are relatively simple to estimate for a given situation, but production losses are more difficult. Methods of dealing with this problem are discussed. Breeding for disease resistance usually requires that either selection candidates, or their relatives, are exposed to the pathogen so that resistance levels can be compared. Parasitic diseases generally create no special ethical problems in a breeding program unless natural challenge levels are insufficient to enable discrimination between hosts in their susceptibility. In the longer term, it is desirable that selection criteria for all major diseases be developed that will be informative in healthy animals. Molecular genetic markers offer promise, but simple genetic markers have so far been as elusive as physiological traits to predict resistance in undiseased animals. In the longer term, useful genetic markers will be found and techniques for combining these with phenotypic information need to be developed. Commercial breeding programs for sheep which include resistance to gastrointestinal roundworms are now operating in Australia and New Zealand, and issues related to breeding in the tropics are discussed.

In merino sheep bred for either increased or decreased resistance to Haemonchus contortus, faecal worm egg counts (FEC) were lower in the resistant line (6,831 vs 17,645 epg, P < 0.01), and circulating eosinophils (EOS) were higher, but not significantly so (3.40 x 10(4) ml(-1) vs 1.40 x 10(4) ml(-1), P = 0.1 1). Another flock was artificially infected with Trichostrongylus colubriformis and significant genetic variation was found in both FEC (heritability 0.40 +/- 0.11) and EOS (0.19 +/- 0.08). In a third flock comprising tropical sheep breeds, a natural challenge with T. colubriformis resulted in significant sire effects on FEC (heritability 0.20 +/- 0.10) but not EOS (heritability inestimable). We conclude that EOS offers no advantage over FEC as it selection criterion for resistance.

Polymorphic bands were detected within the DQB and DRB regions of the ovine major histocompatibility complex by probing TaqI digested DNA from three large sheep half-sib families derived from a highly resistant ram. All animals were phenotypically assessed for Haemonchus contortus resistance by faecal egg counts and associations with RFLP bands and haplotypes were estimated using mixed model, best linear unbiased prediction statistical methods. Although the highly resistant sire was homozygous at the MHC, no significant associations were found between any band or haplotype and faecal egg count.

Peri-parturient worm egg counts were compared in 395 Merino ewes bred for either increased (IRH) or decreased (DRH) resistance to Haemonchus contortus, or from an unselected control flock (CH). Following a 10 month period with no anthelmintic treatment, a rise in egg counts began about 4 weeks before parturition and continued into lactation. At all stages, egg counts were significantly lower in IRH ewes, even those failing to conceive. Before lambing, IRH ewes averaged 263 epg, compared with 1113 epg in CH and 1618 epg in DRH ewes. After 2 weeks of lambing, lactating ewes averaged 1050, 2173 and 3708 epg, respectively, in the three lines. At the end of lambing, egg counts had increased to 1645, 3959 and 4124 epg, respectively. Counts appeared to have peaked in IRH ewes but not in DRH ewes. Ewes with twins had higher counts than those with singles, and ewes suckling lambs had higher counts than those losing lambs. No significant variation was associated with age of ewe (2-7 years). Although there were several nematode species present, the results suggest that, in comparison with DRH ewes, IRH ewes were particularly effective in reducing pasture contamination with H. contortus, both before and during the peri-parturient period.

The leakage of plasma into skin following injection of histamine, bradykinin, activated complement, platelet-activating factor and serotonin was measured in sheep bred for resistance or susceptibility to fleece rot and fly strike. Genetically susceptible sheep had significantly greater plasma leakage to activated complement than genetically resistant sheep, and for all mediators there was a trend for plasma leakage to be greater in susceptible sheep. Within each genotype there was also a tendency for plasma leakage to be positively correlated with fleece rot score. In a flock of sheep of different genetic background not selected for resistance or susceptibility to fleece rot and fly strike, positive phenotypic correlations were also noted between fleece rot and plasma leakage. Plasma leakage provides nutrition for the first instar larvae of Lucilia cuprina, the major cause of primary blowfly strike in Australia. Diminished leakage of plasma following release of endogenous permeability mediators may be one component of the mechanism that confers resistance in animals bred for resistance to fleece rot and fly strike.

A composite population of Haemonchus contortus was established with larvae from seven diverse sources, then maintained in Merino sheep bred to have either increased or decreased resistance to Haemonchus. After five, seven and 14 parasite generations, the two resulting lines of parasites were used to infect sheep from the increased resistance line, an unselected control line and the decreased resistance line. Line of sheep had a highly significant effect on average faecal egg counts 4 and 5 weeks after infection (geometric means 329, 735 and 1490 epg, respectively after the 14th generation), but the two lines of parasites yielded similar egg counts. There was no significant interaction between line of sheep and line of parasite, indicating that the parasite populations had not diverged significantly in their reproductive fitness, as measured by faecal egg count.

Lines of Merino sheep selected for increased (IRH) and decreased (DRH) resistance to Haemonchus contortus were compared with an unselected (CH) line, after approximately four generations of selection. Measurements were recorded on 69 IRH, 47 DRH and 84 CH animals. Following artificial challenge with H. contortus, the IRH line had significantly (P less than 0.001) lower faecal egg counts than the CH and DRH lines (2730, 12,720 and 17,400 epg, respectively). Significant differences (P less than 0.05) were found between all lines in the minimum packed cell volumes during artificial infection (25.7, 22.0 and 20.3%) and in faecal egg counts after natural infection (140, 3590 and 8750 epg). Differences were also recorded (P less than 0.05) following artificial challenge with Trichostrongylus colubriformis (490, 840 and 1340 epg). On a percentage basis, faecal egg counts in the IRH line deviated less from the CH line following artificial infection with T. colubriformis (42%) than with H. contortus (79%). The reverse was true for the DRH line (60 and 37%, respectively). Differences in egg output of this magnitude should have a marked effect on requirements for anthelmintic treatment, rate of development of drug resistance and level of pasture contamination when the lines are grazed separately.