THE EPIDEMIOLOGY OF EQUINE STRONGYLOSIS IN SOUTHERN QUEENSLAND 2. The Survival and Migration of Infective Larvae on Herbage

The seasonal changes in longevity on herbage of the infective larvae of strongylid nematodes of the horse were studied. During the summer months, 1% of the larvae survived on herbage for 2-3 weeks, with 0.2% still viable for a further 2-3 weeks. Equivalent survival periods in winter were 7-11 weeks and over 11 weeks respectively. During spring and autumn, larvae survived for periods varying from 3-8 weeks. On Rhodes grass (Chloris gayana) growing vigorously in the summer of 1976, the majority of larvae remained in the lowest layers of the pasture, within 10 cm of the soil surface. Very few reached the highest fraction of grass sampled, above 40 cm from the soil. More larvae were recovered higher on the pasture in a period when less torrential rain had occurred. It was concluded that the parasitological benefits to be gained from short-term mixed grazing with horses and cattle may be minimal, in view of the tendency of cattle to eat only the upper layers of the pasture initially, with a consequent increase in the number of infective larvae per unit weight of herbage remaining.     

Development and survival ofhaemonchus contortus larvae on pasture in iraq

The survival of Haemonchus contortus infective larvae on pasture and soil was studied over a period of 12 months in the Baghdad area. Infective larvae were found on herbage and soil at all times except in the summer months. During autumn and winter infective larvae in pasture survived for periods of up to 32 weeks. Little larval migration into soil was observed during this study and larvae did not survive for long in the faecal pellets during the summer.     

Observations on the free-living stages of cattle gastrointestinal nematodes

A 4-year study on the free-living stages of cattle gastrointestinal nematodes was conducted to determine (a) the development time from egg to infective larvae (L3) inside the faecal pats, (b) the pasture infectivity levels over time, and (c) the survival of L3 on pasture. Naturally infected calves were allowed to contaminate 16 plots on monthly basis. Weekly monitoring of eggs per gram of faeces (epg) values and faecal cultures from these animals provided data for the contamination patterns and the relative nematode population composition. At the same time, faecal pats were shaped and deposited monthly onto herbage and sampled weekly to determine the development time from egg to L3. Herbage samples were collected fortnightly over a 16-month period after deposition to evaluate the pasture larval infectivity and survival of L3 over time. The development time from egg to L3 was 1-2 weeks in summer, 3-5 weeks in autumn, 4-6 weeks in winter, and 1-4 weeks in spring. The levels of contamination and pasture infectivity showed a clear seasonality during autumn-winter and spring, whilst a high mortality of larvae on pasture occurred in summer. Ostertagia spp., Cooperia spp. and Trichostrongylus spp. were predominant and a survival of L3 on pasture over a 1-year period was recorded in this study.     

Development of free-living stages of equine strongyles in faeces on pasture in a tropical environment

The development of the free-living stages and yields of infective third stage strongyle larvae in faeces from a horse with a mixed natural infection deposited on pasture plots were studied over a 2-year period in a coastal area in tropical north Queensland. Two sets of faecal masses (one exposed to, and the other protected from the action of a natural population of dung beetles) were deposited monthly and after 7 days faecal samples were taken for larval recovery and counts. Hatching and development of the free-living stages occurred in faeces on pasture throughout the year. Development was rapid as infective stages were reached within a week of faecal deposition in all months. Yields of infective larvae were affected by the season and the action of dung beetles on the faecal masses. Highest yields were obtained from both beetle-exposed and protected faeces during winter (June to August) and lowest yields were in spring (September to November). High temperatures in spring and summer resulted in low yields of larvae, however, the dry conditions in spring made this season the most unfavourable period. In autumn and winter the temperatures were never low enough to stop or markedly slow down the rate of development, and allowed the development of large numbers of infective larvae. Dung beetle activity was observed throughout the year, and exposed faeces were usually completely dispersed within 24 h of deposition. This resulted in lower yields of infective larvae from these than from protected faeces. Though larval yields were lower, the actual numbers were still substantial so as to cast doubt on the usefulness of these beetles as biological control agents for equine strongylosis in the dry tropics.     

Seasonal translation of equine strongyle infective larvae to herbage in tropical Australia

Longevity in faeces, migration to and survival on herbage of mixed strongyle infective larvae (approximately 70% cyathostomes: 30% large strongyles) from experimentally deposited horse faeces was studied in the dry tropical region of North Queensland for up to 2 years. Larvae were recovered from faeces deposited during hot dry weather for a maximum of 12 weeks, up to 32 weeks in cool conditions, but less than 8 weeks in hot wet summer. Translation to herbage was mainly limited to the hot wet season (December-March), except when unseasonal winter rainfall of 40-50 mm per month in July and August allowed some additional migration. Survival on pasture was estimated at 2-4 weeks in the summer wet season and 8-12 weeks in the autumn-winter dry season (April-August). Hot dry spring weather (pre-wet season) was the most unfavourable for larval development, migration and survival. Peak counts of up to 60,000 larvae kg-1 dry herbage were recorded. The seasonal nature of pasture contamination allowed the development of rational anthelmintic control programs based on larval ecology.     

Migration of gastrointestinal nematode larvae from cattle faecal pats onto grazable herbage

This study investigated the effect of successive harvests of grazable herbage around cattle faecal pats on the population dynamics of infective gastrointestinal nematode larvae (L(3)). Faecal material, collected from naturally infected calves, was deposited as pats during summer, autumn and winter on three different topographical aspects within a moist, temperate region of New Zealand. Herbage was harvested four times (22-248 days) from around the faecal pats to a height of 2cm in three radial zones (0-20cm, 20-35cm and 35-45cm from the centre of the faecal pat) and L(3) extracted. Harvest date was determined by herbage mass to simulate grazing events. L(3) extracted from herbage were predominantly Cooperia spp. More L(3) were recovered from faeces deposited in summer and autumn, than those deposited during winter. L(3) concentration on herbage was highest (P<0.001) in the zone nearest the pat for all except the fourth harvest. Mean concentrations of L(3) on herbage were 11,447, 3154, 337 and 102 L(3)/kg dry matter herbage, for the four successive harvests, respectively. Microclimate differences as affected by aspect had a marked effect on herbage growth, but did not significantly affect L(3) concentration on herbage. In this study, L(3) remained aggregated close to the faecal pats they emerged from even after two successive harvests and significant rainfall. Successive harvests simulated the effect of repeated grazing events by a non-infective stock class. Two such grazings and the associated time, reduced L(3) presence on grazable herbage to <3% of the original population. Grazing strategies to generate clean pasture for vulnerable cattle are discussed in relation to these results.     

Development and survival of infective larvae of gastrointestinal nematodes of cattle on pasture in central Kenya

On a series of pasture plots, 2 kg pats of bovine faeces containing known numbers of strongylid (Haemonchus, Cooperia, Oesophagostomum and Trichostrongylus) eggs were deposited at intervals of 4 weeks from July 1995 to June 1996. The plots were sampled every 2 weeks after contamination and infective larvae were identified and counted. Larvae of all the genera developed throughout the year, but the pats exposed during the rainy season yielded more abundant larvae on the herbage. Irrespective of the season of deposition of the pats, larvae were found in larger numbers from 2 to 6 weeks after deposition and generally declined to below detectable levels within 12 to 16 weeks of contamination. The comparatively short survival times noted in this experiment may present opportunities for manipulation of the population dynamics of the gastrointestinal nematodes in the tropical environment of Kenya.     

Epidemiology of Nematodirus species infections of sheep in a subarctic climate: development and persistence of larvae on herbage

As part of a study on the epidemiology of Nematodirus species of sheep in subarctic Greenland, the development and persistence of eggs and larvae were investigated by experimentally contaminating plots of pasture with infected faeces and by placing tubes containing a suspension of eggs on to or into the soil. Despite low ambient temperatures, infective larvae appeared within a month during the summer. The greatest numbers of larvae were recovered from herbage in August and September. Eggs did not develop synchronously as development beyond the morula stage could be delayed for up to two years. Larvae were found on herbage for up to 37 months after faecal deposition. In the sheep rearing area of Greenland, therefore, Nematodirus species larvae can be present on herbage throughout the whole summer but peak numbers occur late in the grazing season.     

Effect of different times of administration of the nematophagous fungus Duddingtonia flagrans on the transmission of ovine parasitic nematodes on pasture--a plot study

Investigations were made into the timing of administration of Duddingtonia flagrans as a biological control agent against ovine parasitic nematodes including stongylid and Nematodirus spp. Faeces from 3-4 months old male lambs were deposited onto pasture plots that had never been grazed by sheep. The trial was conducted over two consecutive years (1998 and 1999). For both years, the following three plot types were involved: Sim plots had faeces containing nematode eggs and Duddingtonia flagrans spores deposited simultaneously; Post plots had faeces containing nematode eggs followed 2 weeks later by faeces containing D. flagrans spores alone; Control plots had faeces containing only nematode eggs; Prior plots (included in 1999) had faeces containing D. flagrans spores alone followed 2 weeks later by faeces containing nematode eggs. In each year, two deposition periods were involved: July and August in 1998 and June and July in 1999. During the first year pasture samples were collected at 2, 4, 6, 8 and 12 weeks after initial deposition. In 1999, additional samples were collected at 10, 16 and 20 weeks. Larvae were extracted from the pasture samples and counts performed to estimate the number and species of infective third-stage (L(3), larvae) present. The number of third-stage strongylid larvae on pasture was significantly lower on Sim plots compared to the remaining plot types for both years at all deposition times (P<0.001). This was also the case for the number of Nematodirus infective larvae in August deposition plots in 1998 (P<0. 02). There was no significant difference between treatments in both deposition times in 1999 and July deposition plots in 1998 for the Nematodirus data. These results suggest that D. flagrans, if deposited at the same time as parasite eggs prevents transmission of third-stage larvae from the faecal deposit onto pasture, including occasionally Nematodirus species, but does not have an effect on third-stage parasitic nematode larvae in the surrounding soil.     

Epidemiology of bovine dictyocaulosis in Denmark

A field experiment was conducted over two grazing seaons with calves on a permanent pasture in order to follow the pattern of infection with Dictyocaulus viviparus. Infective larvae persisted during the first, but not during the second, winter of observation. By means of the agar-bile herbage technique, a moderate first peak of infection was demonstrated in the pasture 2–3 weeks before the appearance of respiratory signs in the calves. Fluctuations in faecal larval output were reflected in the herbage contamination with infective larvae close to faecal pats. This, as well as the horizontal dispersion of larvae in the pasture, took place in less than a week. The proportion of lungworm larvae recovered away from faeces was low during a period of dry and hot weather while herbage sampling at two-hour intervals during two days showed an increase in herbage contamination with lungworm larvae, but not with trichostrongyle larvae between 10 a.m. and 12 noon.The infectivity of the pasture was monitored by tracer calves and compared with the results of the pasture sampling. The general course of the infection in the calves and in the pasture was the results of interaction between them. In addition, the pasture infection was influenced by climate and the infection in the calves by the development of immunity. The course of infection in individuals appeared to have an influence on the general course of the infection through the contamination of the pasture.     

Epidemiological investigations of trichostrongylid infections in young cattle in different parts of Norway

The occurrence of trichostrongylid nematodes in young cattle was studied in relation to climate and geographical location, by examination of herbage and faecal samples from 8 farms in 7 different regions in Norway, during 3 consecutive summers. Species of Ostertagia and Cooperia were dominant. Infective larvae, especially those of Ostertagia, were able to survive the winter in the pasture to a great extent in all the areas studied. The spring larval counts varied considerably from place to place, and from year to year, but were often at fairly high levels. This confirms previous findings that the larvae which overwinter on the pasture may be a major cause of losses. Overt clinical disease, however, was only seen at 1 farm during this investigation.The large number of larvae often seen in the autumn seems to be of less significance as a cause of disease, due to the late appearance, but is of major importance as a source of the larvae which survive the winter in the pasture and infect young cattle the following spring.There was no obvious correlation between pasture larval counts and the climate, except that there seemed to be higher counts in the warmer parts of the country, where the grazing season is also longer. The only place were both faecal egg counts and larval counts in the herbage were consistently low, was in the very far north.     

The development of Trichostrongylus colubriformis larvae on a range of herbage species or on plots of differing topographical aspect

Five "contaminations", where faeces containing Trichostrongylus colubriformis eggs were deposited on pasture and serially recovered, were used to compare the rate of decline of faecal mass and larval development. In the first three contaminations, faeces from a common source were deposited on swards of browntop (Agrostis capillaris cv Grasslands Muster), ryegrass (Lolium perenne cv Grassland Nui), white clover (Trifolium pratense cv Grassland Tahora), or onto bare ground in the late spring, summer or autumn. The last two contaminations were done on the north facing aspect or south facing aspect of hill country pastures in summer and autumn. Number of free-living nematodes (first- and second-stage larvae (L(1) and L(2)) and soil dwelling nematodes) and third stage larvae (L(3)) recovered from faeces were counted. In spring there was a significant (P<0.01) effect of sward type on the mass of faeces remaining, with greatest mass remaining on browntop and ryegrass 28 days later, and less on bare ground and white clover. In summer there were more (P<0.05) faeces remaining on browntop than on other herbages which had little faeces remaining and which did not differ one from another. In autumn there was a rapid decline in faecal mass. All faeces were gone from white clover and ryegrass swards by day 10 and from browntop and bare ground by day 14. The number of free-living nematodes did not differ markedly between seasons, ranging from 5 to 8.5% of eggs deposited. The number of L(3) recovered was low in spring ( approximately 0.4% of eggs deposited) and did not differ between swards. In summer, more (P<0.05) L(3) were recovered from faeces deposited on swards of ryegrass and white clover than from bare ground or browntop. Most L(3) were recovered from days 7 to 14 ( approximately 1.3% of eggs deposited). In the autumn, low numbers of L(3) were recovered from browntop on day 3 and ryegrass on day 7 (0.2% of eggs deposited) with virtually no L(3) recovered from faeces placed on white clover or bare ground. There were significant (P<0.001) effects of aspect on the amount of faecal mass remaining in both summer and autumn with less faeces remaining on the south facing aspect than on the north. This was particularly evident during the summer when virtually all of the faeces were intact on the north facing aspect but only 40% was remaining on the south on day 28. In the autumn, while faeces were completely gone from both aspects by day 28 but there were less (P<0.05) faeces remaining on the south facing aspect from days 3 to 18 than from the north. There was no aspect effect in either season on the number of free-living nematodes recovered which averaged 8-11% of eggs deposited. In both seasons a greater number of L(3) were recovered from faeces on the south facing aspect than on the north, particularly 3-10 days after faecal deposition. In summer the rise in L(3) recovered in faeces was more rapid on south facing aspect than on the north but both attained a maximum level of approximately 4% of eggs deposited. In autumn on day 3 there was a rapid rise on south facing aspect to approximately 21% of eggs deposited followed by a gradual decline on day 10 while on the north facing aspect numbers of L(3) recovered only attained 10% of eggs deposited.     

Studies on the epidemiology of bovine parasitic bronchitis.

In the West of Scotland the epidemiology of parasitic bronchitis in grazing calves was studied over a two year period with the aid of tracer calves and herbage examinations for Dictyocaulus viviparus larvae. The observations of both years emphasised the importance of overwintered lungworm larvae as a source of disease. In the first year it was shown that the ingestion and development of these overwintered larvae were, by themselves, directly responsible for severe morbidity, high faecal larval counts and deaths. In the second year it was shown that pasture ungrazed during the winter and spring and from which a hay crop was removed in mid-summer was still capable of producing clinical parasitic bronchitis in susceptible calves within three to four weeks of their introduction in later summer. In both years there was some evidence that the outbreaks appeared to be associated with the sudden availability of infective larvae on the herbage. The possibility that such larvae may have survived for many months in the soil is discussed. Despite the heavy challenge with lungworm larvae experienced by the grazing calves in the first year those vaccinated with lungworm vaccine survived, their clinical signs were mild and of short duration and their faecal larval output was greatly reduced.     

Development and spatial distribution of the free-living stages of Teladorsagia circumcincta and Trichostrongylus colubriformis on pasture: A pilot study

Abstract

AIMS: To measure the development of Teladorsagia (=Ostertagia) circumcincta and Trichostrongylus colubriformis eggs to third-stage infective larvae (L3) at different times of the year. Also, to measure the spatial distribution of L3 across herbage, soil and faeces, in order to assess whether spatial issues could be important in larval dynamics on pasture.

METHODS: Field plots were contaminated with sheep faeces containing approximately 20,000 eggs of each of T. circumcincta and T. colubriformis on five separate occasions, viz 01 December 1996 (summer), 18 March 1998 (autumn), 17 June 1998 (winter), 15 October 1998 (spring), and 23 July 1999 (winter). Replicate plots (n=10) were harvested at intervals for up to 12 months after deposition of faeces, and the number and distribution of L3 were measured. Larvae were sampled from faeces (where these remained), herbage, and three soil zones to a depth of 145 mm.

RESULTS: There were large differences between contamination dates in the percentage of eggs that developed to L3. For both species the highest percentage development was for eggs deposited in December (7.8% and 25.9% for T. circumcincta and T. colubriformis, respectively) and the lowest for June (0.4% and 0.03% T. circumcincta and T. colubriformis, respectively). Development in winter was often delayed, and this was always associated with a low yield of larvae, probably due to compounding mortalities associated with long periods of exposure to low temperatures.

The relative distribution of L3 present on herbage, in faeces or in the soil varied between sampling times. However, overall the most L3 were recovered from soil (74% and 66% for T.circumcincta and T. colubriformis, respectively, averaged over all samples), and the lowest recoveries were from the herbage.

CONCLUSIONS: Although the data are limited, the results indicated that the highest percentage of eggs developed to infective larvae in summer and only minimal development occurred in winter. The data do not support the view that substantial contamination of pastures with sheep parasites occurs over winter. Large numbers of larvae were recovered from soil, which indicates that, assuming they can subsequently migrate onto herbage, soil is a potentially important reservoir ofinfective larvae in New Zealand. Therefore, the spatial distribution of L3 on pasture may affect both the dynamics and transmission of parasite populations. Further work on both these issues is warranted.     

The epidemiology of Dictyocaulus filaria in north east England.

The seasonal pattern of Dictyocaulus filaria infection in four flocks of sheep under field conditions was studied by faecal examination for larvae. The prevalence of infection in lambs was low in spring and summer but increased in late autumn or winter, then fell again to a low level by May. However, most lambs became infected at some time during their first year of life. In ewes the prevalence of infection was generally low, but rose during winter and spring. Seasonal pattern and intensity of infection were influenced by weather conditions and the persistence of infection from the previous year, moist summer conditions giving rise to a higher level of infection in autumn and winter than a dry summer. The source of infection may be the ewes or yearlings, or infective larvae surviving over the winter on pasture. Autoinfection occurred, some lambs becoming infected with at least three generations of the parasite, but at the levels of infection recorded clinical disease was rare.     

Seasonal occurrence of infective nematode larvae in California Sierra foothill pastures grazed by cattle

Worm-free Holstein tracer calves were used to assess the seasonal variation in number and course of development of nematode parasites of cattle on an irrigated pasture located in the California Sierra foothills. The most common genera of nematodes found in the abomasum and small intestine were Ostertagia and Cooperia, respectively. The infective Ostertagia larvae on herbage were most numerous in early spring and lowest in summer months. Arrested development of Ostertagia occurred in mid-to-late spring. The percentage of the population of Ostertagia found to be arrested larvae was constant (73% to 95%) over 4 years, even though the total number of nematodes in the genus was markedly different. Levels of infection with Ostertagia and Cooperia were found to correlate with pasture contamination occurring in the preceding fall and concurrent winter months. In the cecum and large intestine, Oesophagostomum venulosum was found to be the most common nematode. It is suggested that this species may replace Os radiatum as the cattle nodular worm in regions where climate is similar to that of the southwestern coast of North America. Evidence of seasonal hypobiotic development of Oe venulosum was found. Other genera of nematodes observed in tracer calves were Trichostrongylus, Haemonchus, Bunostomum, Trichuris, and Dictyocaulus.     

Observations on the output of nematode eggs in faeces and on the subsequent pasture infestation with third stage larvae produced by a herd of farmed red deer (Cervus elaphus).

More than 2900 faecal samples and 200 herbage samples were examined from January 1986 to January 1989 during a study of the host/parasite relationships of the gastrointestinal nematodes of a herd of farmed red deer in Surrey. The pattern of faecal egg output from the hinds appeared to conform to a tri-modal distribution with peaks during spring, the periparturient period and late summer. Third stage larvae were present on the herbage during every month. The new generation appeared in June but levels of larvae remained low until the autumn. The implications of these findings are discussed in relation to the farming of red deer.     

Factors affecting the survival and migration of the free-living stages of gastrointestinal nematode parasites of cattle in central queensland

Faecal pats containing parasitic nematode eggs were deposited monthly on worm-free pasture, from mid-1975 to early in 1979, near Rockhampton in central Queensland. Pasture samples were collected monthly from beside these pats and the number of infective larvae on the samples was counted.

Cooperia spp. were the most numerous larvae on pasture all year round and Haemonchus placei were commonly present in low numbers. Small numbers of Oesophagostonum radiatum larvae were found, mostly during summer.

Dung beetle activity and rainfall influenced larval populations on pasture, but temperature did not. Beetles were not active in winter, and pats deposited in spring, summer and autumn when beetles were active yielded only 42, 44 and 26%, respectively, as many larvae per 1000 eggs deposited as winter pats. Pats in which beetle activity was minimal (feeding only), moderate and intense (complete destruction), yielded 43, 10 and 6%, respectively, as many larvae per 1000 eggs as intact pats.

Larval densities on pasture were highest after the first saturating rains during the spring-summer period and most of these larvae migrated from unattacked pats deposited in winter. Beetle numbers and activity increased with the summer rains and so few larvae were available to migrate onto pasture during late summer and autumn when the highest falls of rain were recorded. The regression of larval recovery on rainfall was positive and statistically significant when data collected soon after these very heavy rainfall periods were omitted from the analysis.

In 1977, drought-breaking rains increased the normal larval density on pasture 10-fold because larvae in pats deposited in the last 4 months of the drought migrated onto pasture immediately after the rains.

This work suggests that in summer rainfall areas where dung beetles are active, helminth control may be achieved by reducing the worm egg output from cattle during the winter.     

The origin and overwintering survival of the free living stages of cattle parasites in Sweden

During the 1997 Swedish grazing season, faeces were collected every 3 weeks on 7 occasions from young grazing cattle with moderate nematode parasite infections. From this source 12, 400 g dung pats were set up on each sampling occasion on a specially designated area of pasture. Half of these pats were placed on pasture where it was aimed to prevent snow cover during the subsequent winter. During the grazing season, herbage growth was kept at reasonably uniform height by clipping and the dung pats were protected from destruction by animals and birds. At the time of animal turn-out the following year (7th April 1998), it was observed that all dung pats had disappeared. Assessments of the survival of infective larvae, both on pasture and in soil, were made in a circular area encompassing the location of each pat. These sampling procedures were completed within a 3 week period. All faecal deposits yielded infective larvae at turn-out the following year, with proportionally greater numbers developing from nematode eggs deposited in cattle dung during the mid third of the previous grazing season. The surface layer of soil was found to be an important reservoir for infective larvae, with numbers recovered being approximately half those found in the overlying pasture samples. No significant differences were found between the normal pasture and snow excluded pasture in the number of infective larvae recovered from both pasture and soil samples. The epidemiological consequences of these findings are discussed.     

SEASONAL AVAILABILITY OF NEMATODE LARVAE ON PASTURES GRAZED BY CATTLE IN NEW SOUTH WALES

Trends in the numbers of infective nematode larvae on pasture plots contaminated by cattle at different seasons of the year were defined in 3 different climatic regions. The main nematodes were Ostertagia ostertagi, Trichostrongylus spp, Haemonchus spp and Cooperia spp.
On the North Coast of New South Wales with a sub-tropical climate, the numbers of infective larvae of all 4 nematodes rose rapidly to peak levels soon after each seasonal period of contamination began, then fell quickly within a few months. On the Central Coast of New South Wales, the trends were similar to those on the North Coast, except that the larvae persisted on the pasture for a much longer time. On the Northern Tablelands of New South Wales, where temperatures were much colder than on the coast, larval development was slower and major peaks of larval availability did not occur until early spring. These different seasonal trends in each region were considered to be related to the climatic differences between the regions.
On pastures which were contaminated continuously, larval numbers reached maximum levels in mid-winter on the Central Coast and in early spring on the Northern Tablelands. It was concluded that the majority of these larvae were derived from the contamination of pastures in autumn and winter. Subsequently in summer, a rapid dying out of larvae was observed in all the regions, probably due to the effect of hotter weather.
The studies suggest that a reduction in the contamination of pasture with nematode eggs in autumn and winter could result in pastures carrying fewer larvae and thus form the basis of effective worm control programs for cattle.