Mechanisms of survival of nematode parasites with emphasis on hypobiosis

Mechanisms involved in parasitic nematode survival must be considered with reference to their host and environmental interrelationships since these interrelationhips ultimately influence any parasite adaptations aimed at survival.The most important of the potential environmental constraints are climatic, particularly temperature and humidity, and these can drastically influence larval development and survival.One of the major host factors influencing successful parasite survival is the availability of suitable (susceptible) new hosts to the infective stages of the parasite at the appropriate time for transmission to be achieved. Other host factors that influence parasite survival are those that affect the entrance, establishment and reproduction of the parasite within its new host; mainly problems of acclimatization to a parasitic way of life as well as the countering or adaptation to a variety of host resistance factors, both molecular and cellular, by the parasite.Finally in order for life cycles to be completed, the parasite must evolve means whereby its larval forms can leave the host so that eventual transmission to a new host can be accomplished.In this paper a number of adaptations which enable the parasite to overcome these constraints are discussed. These include such things as larval resistance to environmental effects, the utilization of intermediate hosts or vectors for transmission, seasonally-increased fecundity rates, anti-host immunity stratagems and hypobiosis.This latter phenomenon, hypobiosis or prolonged but temporarily arrested larval development, represents one of the most useful of life cycle adaptations to ensure parasite adaption enables the parasite to synchronize its life cycle to changing environmental or survival and appears to be widespread among parasitic nematodes. Among its benefits, this host conditions. It can thus be of major importance in ensuring survival of the parasite during periods of environmental adversity when conditions for transmission are poor and survival of free-living forms may be minimal. It also enables the parasite to have available large numbers of infective forms at points in the host reproductive cycle which coincide with the production of the susceptible neonates thereby greatly facilitating transmission. Additionally, with certain species of nematodes, occurring as it does at times of the year when the numbers of infective stages may be high while host resources may be limited, oscillations in parasite biomass can be avoided. It thus serves as a highly adaptive mechanism for regulating populations of adult worms, lessening stress on the host and favoring parasite survival as a result.     

Hypobiosis and the periparturient rise in sheep

From a parasite viewpoint, therefore, hypobiosis is a most useful life cycle adaptation to ensure persistence and has been reported in most of the important nematode species parasitizing sheep in North America. It facilitates the synchronization of the nematode life cycle to changing host and environmental conditions. It also enables the parasite to have available large numbers of infective forms at points in the host life cycle that coincide with the presence of susceptible neonates, thus ensuring transmission. The accompanying periparturient rise seen with many of these species following development of hypobiotic larvae has very important implications for the planning of control procedures. Because it appears to be the major source of pasture contamination early in the grazing season, it is the prime source of infection for the lambs. These lambs can in turn serve to augment pasture contamination, leading to very heavy levels of pasture infectivity in late summer and fall. By scheduling anthelmintic treatments so as to suppress this rise in egg counts, pasture contamination can be kept to a minimum. More effective, less frequent dosing with anthelmintics is needed.     

The effect of Plasmodium gallinaceum on a challenge infection with Ascaridia galli in chickens

The effect of a primary infection with the haemoparasite Plasmodium gallinaceum on the establishment of a challenge infection with the nematode Ascaridia galli in chickens was studied. Four groups were infected as follows. Group 1: inoculated intravenously with 10(6) P. gallinaceum-infected erythrocytes on day 0; group 2: orally infected with 500 embryonated A. galli eggs on day 10; group 3: infected with P. gallinaceum on day 0 and A. galli on day 10; and group 3: non-infected control birds. The results of this investigation demonstrates that a primary infection with P. gallinaceum in chickens alters the course of a subsequent infection with A. galli. Thus, an antagonistic effect was seen in which the malaria infection caused a significant reduction on the establishment of the nematode in concurrently infected animals.     

文昌鸡生态放养过程中驱虫程序的研究

采用虫卵漂浮法、麦克马斯特氏虫卵计数法对文昌鸡生态放养期间感染寄生虫的种类与粪便中虫卵数量进行监测。结果表明,文昌鸡生态放养感染的寄生线虫主要为鸡蛔虫和几种异刺线虫。自然条件下生态放养35 d后,每克粪便中寄生虫虫卵数（eggs per gram, EPG）达到了700,呈显著增长趋势。抽样宰杀30只鸡,检出18只感染鸡蛔虫成虫,感染率为60％,平均感染强度为8条/只。因此,建议整个生态放养期间驱虫3～4次为宜,首次驱虫时间为放养后25～30 d。     

Development of resistance to Trichostrongylus colubriformis in goats

A high level of host resistance to Trichostrongylus colubriformis was established in a group of Saanen goats infected weekly with 10,000 infective larvae for 10 weeks, treated with an anthelmintic and then challenged twice with 50,000 larvae, with a further anthelmintic treatment 28 days after the first challenge. A second group of goats exposed only to the two challenge infections developed worm burdens intermediate between, and significantly different from, the first group and a third control group given only the second challenge infection. Nematode fecundity (eggs per female) and male:female ratios were also significantly decreased in the first two groups. Faecal egg counts and worm counts were highly correlated and this relationship was dominated by the influence of the number of eggs/female nematode. The goats were approximately 9 months old at the start of the experiment.     

Effect of the consumption of Lysiloma latisiliquum on the larval establishment of gastrointestinal nematodes in goats

The consumption of tannin-rich (TR) forages has been associated with negative effects against gastrointestinal nematodes and with an improved host resilience. It has been hypothesized that tannins affect the capacity of infective larvae to establish in the mucosae of the host. In this study, we aimed at testing this hypothesis using Lysiloma latisiliquum, a tropical TR tree. The objectives were: (i) to evaluate the effect of the consumption of L. latisiliquum on the establishment of nematode third-stage larvae (L3) in goats; (ii) to define the role of tannins in these effects in vivo by using an inhibitor (polyethylene glycol, PEG); and (iii) to examine a possible indirect effect of tannins on the inflammatory response in the digestive mucosa. Eighteen Criollo goats composed three experimental groups. The control group received fresh leaves of Brosimum alicastrum, a plant with a low level of tannins. Two groups received L. latisiliquum leaves either with (L.L.+PEG) or without (L.L.) daily addition of 25g PEG. After a 7-day adaptation period, each goat was infected with both Haemonchus contortus and Trichostrongylus colubriformis (3000 L3 per species). The goats were slaughtered 5 days after infection and worm counts and histological analyses were performed. No difference in the voluntary feed intake of foliage was observed between the 3 groups. The consumption of L. latisiliquum significantly reduced the larval establishment of both nematode species compared to the control (P<0.01). For both worm species, the effects were totally alleviated with PEG (L.L.+PEG group), suggesting a major role of tannins in the observed effects. Only minor differences in the mucosal cellular response were observed between the 3 groups. These results confirm that the consumption of TR plants reduces the establishment of nematode larvae in the host and that a direct effect is principally involved.     

Infection of Japanese quail with Toxocara canis larvae and establishment of patent infection in pups

Migration and distribution of Toxocara canis larvae in the tissues of Japanese quail, infected orally with 5 X 10(3) infective eggs, and the establishment of patent infection from the quails to the definitive host, were studied. Larval yield at necropsy from various tissues and organs of quail varied from 4.72 to 7.54% of the infective eggs inoculated within the period 1-60 days post-infection (PI). The total number of larvae recovered on different days showed a gradual increase. The percent inoculum recovered at necropsy was highest on Day 60. Most of the larvae were found in the liver throughout the period and only a few migrated to other tissues, such as lung, heart, muscle and brain. The establishment of patent infections in the definitive host was studied by feeding the 15-day infected livers of Japanese quail (400 larvae) to Toxocara-free pups. Eggs first appeared in the faeces 38 days post-infection, the mean worm burden at necropsy was 87 and the percentage of infection established was 21.75%. Thus the role of Japanese quail as a paratenic host of T. canis is established.     

Duration of the persistent activity of moxidectin against Haemonchus contortus in sheep

SUMMARY Weaned lambs were infected with Haemonchus contortus 35,28,21,14 and 7 days after treatment with moxidectin at 0.2 mg/kg and 35 and 14 days after treatment with ivermectin at the same dose rate. Worm counts 14 days after infection showed that moxidectin prevented the establishment of over 99% of infective larvae for 28 days and reduced the establishment rate at 35 days by 96%, relative to ivermectin. There was no difference in the protective efficacy of ivermectin at 14 or 35 days. The persistence of moxidectin is likely to provide advantages in nematode control, particularly when used as a strategic early summer treatment or as a pre-lambing treatment to ewes. Implications of the persistent activity of moxidectin for the development of resistance during the decay phase are discussed.     

Effects of the repeated distribution of sainfoin hay on the resistance and the resilience of goats naturally infected with gastrointestinal nematodes

Due to the high prevalence of anthelmintic resistance in goats, the need to explore novel approaches to control nematodes and to reduce the exclusive reliance on chemotherapy is strongly demanded in this host species. In sheep, several studies have shown that the consumption of tannin-rich legume forages was associated with positive effects on host resilience and resistance to parasite infection. In goats, studies on such interactions between tanniferous plants and nematode infections remain few. The objectives of the current study were to examine under natural conditions the effects of consumption of sainfoin hay by goats on the parasite populations and on host resilience. Eighteen adult cull goats naturally infected with Haemonchus contortus, Teladorsagia circumcincta and Trichostrongylus colubriformis were used in the study. At the start of the assay, the goats were allocated into two groups, balanced according to weight and the levels of egg excretion. The two groups grazed separate pastures for 3 months with similar stocking rates. Goats from group S received each month indoors, for 7 days, sainfoin hay and control goats (group C) received hay of ryegrass. The diets in both groups were made isoenergetic and isoproteic and the refusals measured. Individual parasitological and pathophysiological measurements were performed fortnightly in order to compare host resistance and resilience. At the end of the study, five goats per group were necropsied. The distribution of sainfoin was associated with: (1) a higher consumption of hay; (2) significant, lower levels of nematode egg excretion which was associated with a decrease in worm fertility but no change in worm population; however, the number of intestinal worms was reduced by 50% in group S; (3) a better host resilience. In particular, after 2 months of grazing, two control goats died and half of the remaining animals needed to be treated whereas this was not the case in group S. These differences were related to significant changes in pepsinogen and phosphate values (PCV) but not in pepsinogen and phosphate concentrations. These results demonstrate that a repeated distribution of sainfoin hay to grazing goats might be beneficial in regard of pasture contamination and host resilience. They suggest that administration of sainfoin hay might represent a valuable alternative and adjunct to reduce nematode infections in dairy goat flock.     

Larval viability and serological response in horses with long-term Trichinella spiralis infection

The horse is considered an aberrant host for the nematode parasite Trichinella spiralis, and many aspects of the biology and epidemiology of Trichinella infection in the horse are poorly understood. It has been reported that experimentally-infected horses produce a transient serological response to infection and that muscle larvae are cleared more rapidly than in parasite-adapted hosts such as the pig and humans. However, limited numbers of animals have been studied, and both the longevity of larvae in horse musculature and the immune response to Trichinella larvae remain unclear. In this study, we infected 35 horses with 1000, 5000, or 10,000 T. spiralis muscle larvae and followed the course of infection for 1 year, assessing larval burdens in selected muscles, the condition and infectivity of recovered larvae, and the serological response of infected horses. The results demonstrated that T. spiralis establishes infection in horses in a dose dependent manner. Anti-Trichinella IgG antibodies peaked between weeks 6-10 post-inoculation. Viable, infective larvae persisted in horse musculature for the duration of the study (12 months), and exhibited no apparent reduction in muscle burdens over this period. Encapsulated larvae showed no obvious signs of degeneration in histological sections. Larval capsules were surrounded by infiltrates consisting of mature plasma cells and eosinophils. Macrophages were notably absent. Given the lack of a detectable serological response by 26 weeks p.i. and the persistence of infective muscle larvae for at least 1 year, parasite recovery methods are currently the only suitable detection assays for both meat inspection and epidemiological studies of Trichinella infection in the horse.     

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.     

The occurrence of Contracaecum sp. larvae (Nematoda: Anisakidae) in the catfish Clarias gariepinus (Burchell) from Lake Chivero, Zimbabwe

Clarias gariepinus were collected from Lake Chivero, Zimbabwe, and examined for nematode parasites from November 2000 to May 2002. Of the 202 specimens collected, 42.6% were infected with third-stage larvae of Contracaecum sp. in the body cavity. The intensity of the infection was 1-7 worms per fish (mean intensity = 2.2). Seasonal variation in the prevalence of the parasite was not obvious and there was no significant difference in the prevalence of infection between males and females (chi 2 = 2.228; P > 0.05). No significant relationship between host size and prevalence was established. There was also no significant relationship between intensity and the body condition factor (r = 0.11; P > 0.05). The low parasite prevalence may have been caused by the disruption of the infection cycle since piscivorous birds, which are the final hosts of the parasite, do not feed on C. garieplnus in Lake Chivero.     

Further study of Codiostomum struthionis (Horst, 1885) Railliet and Henry, 1911 (Nematoda, Strongylidae) parasite of ostriches (Struthio camelus Linnaeus, 1758) (Aves, Struthioniformes)

Codiostomum struthionis is a nematode parasite of the ostrich caecum. Little is known about its pathology, being considered by many authors as a non-pathogenic parasite. Infections by C. struthionis are sometimes overlooked because its eggs are indistinguishable from another ostrich nematode, Libyostrongylus spp. Fecal cultures and infective larvae identification are necessary for proper identification. The aim of this study is to provide improved morphological characterization of adults and infective larvae of C. struthionis. Ten caeca of adult ostriches were collected and washed in 0.09% saline solution. Male and female nematodes were collected and quantified separately. Nematodes were fixed in A.F.A. for optical microscopy or fixed in Karnovsky solution for scanning electron microscopy. To obtain infective larvae, fecal samples were collected at sites of high concentration of parasites in the caeca and fecal cultured. The resultant larvae were identified and measured with light microscope at 400x. Nine of the 10 slaughtered ostriches were parasitized by C. struthionis. All nematodes were found in the distal third of the caeca. A total of 566 parasites were recovered (234 males and 332 females). All the cultured larvae had characteristics of C. struthionis (rounded cephalic region with a flat extremity, an acute larvae tail termination and a long and filamentous sheath tail). All the adult parasites were characterized as C. struthionis. Through the analysis of the infective larvae it was determined that the morphology of the larvae tail was the best trait to use in the distinction of this species (live bird diagnosis).     

Changes in food intake and circulating leptin due to gastrointestinal parasitism in lambs of two breeds

A reduction in food intake is a prominent feature of many infectious diseases. However, the underlying mechanisms of parasite-induced anorexia in sheep are poorly understood. Here, we tested the hypotheses (a) that the degree of parasite-induced anorexia in lambs is influenced by their growth potential and (b) that nematode infection results in elevated plasma leptin concentration in lambs. The hypotheses were tested with Suffolk x Greyface (S) and Scottish Black-face (B) lambs that are known to differ in their growth potential (S lambs are of greater growth potential than B lambs). During a primary parasite infection, 24 out of 48 lambs per breed were trickle-infected with 7,000 infective Teladorsagia circumcincta larvae per day, 3 d/wk, for a period of 12 wk (experiment I). The lambs were then dewormed, and after a 2-wk interval, half of the 24 lambs per breed that were previously infected were reinfected for another 12 wk with the same parasite and dose as used in the primary infection (experiment II). In both experiments, infected lambs were fed grass pellets for ad libitum intake, whereas noninfected lambs were fed grass pellets for either ad libitum or restricted intakes. The S lambs were more susceptible than B lambs to nematode infection, as judged from the differences in fecal egg counts (P = 0.007). Parasitized lambs of the more susceptible breed (S) showed anorexia [i.e., a decrease in intake of 13% compared with uninfected controls (P = 0.01)], whereas no significant reduction in food intake was observed in lambs of the more resistant breed (B). Reexposure to nematode infection of previously infected animals tended to result in renewed anorexia in S lambs but not in B lambs (P = 0.08) in a similar extent as during primary infection. Plasma leptin concentrations did not differ between ad libitum-fed infected and control lambs but were greater in infected than in noninfected lambs at a similar level of food intake during both the primary (P = 0.02) and the secondary parasitic infection (P = 0.004) in both breeds. The results show that leptin may be involved in the response of lambs to infection but that it is unlikely that leptin alone is responsible for the parasite-induced anorexia in lambs.     

Genetic analysis of the potential role of IgA and IgE responses against Haemonchus contortus in parasite resistance of Creole goats

Small ruminants are affected by gastrointestinal nematode (GIN) infection. A promising alternative strategy for control of GIN infection is to increase the level of resistance in the population by taking advantage of the host's immune response. Immunoglobulin A (IgA) and E (IgE) are known to be involved in immune response to GIN. The aim of this study was thus to investigate genetic parameters of IgA and IgE responses against Haemonchus contortus in Creole kids naturally challenged at pasture and to determine the relationship with other resistance criteria such as faecal egg counts, packed-cell volume, eosinophil counts and bodyweight. Variance and covariance components for genetic and residuals effects for each trait were estimated on 3862 males at 11 months of age. Heritability estimates for IgA and IgE ranged between 0.15 and 0.57. Strong positive genetic correlations were observed between either IgE or IgA responses against L3 and adult excretory/secretory products (ESP) antigens of H. contortus, suggesting that the humoral immune response is not specific to the life cycle stage of the parasite suggesting that there is substantial cross recognition between the different parasite antigens. Heritability estimates for faecal egg count (FEC), packed-cell volume (PCV) and bodyweight (BW) were in accordance with previous results in Creole kids. Blood eosinophil counts were found moderately heritable and negatively correlated with FEC, suggesting that this cell population plays a role in resistance to nematode parasite infection in Creole goats. IgA response was positively correlated to FEC, in contrast with the negative correlation between IgE against L3 of H. contortus and FEC. In Creole goats, IgA response against L3 or ESP of H. contortus would rather be associated with the worm burden than an immune protective response. The immune response involving activity of IgE against L3 of H. contortus may be one important pathway for development of resistance to gastrointestinal nematode infections in Creole goats.     

雏鸡感染堆型艾美耳球虫及体内一氧化氮的生成

本研究利用Griess反应监测了雏鸡感染堆型艾美耳球虫(Eimeriaacervulina)后血浆NO     

Potential for dissemination of the nonnative salmonid parasite Myxobolus cerebralis in Alaska

Myxobolus cerebralis, the myxozoan parasite responsible for whirling disease in salmonids, was first introduced into the United States in 1958 and has since spread across the country, causing severe declines in wild trout populations in the intermountain western United States. The recent detection of the parasite in Alaska is further evidence of the species' capability to invade and colonize new habitat. This study qualitatively assesses the risk of further spread and establishment of M. cerebralis in Alaska. We examine four potential routes of dissemination: human movement of fish, natural dispersal by salmonid predators and straying salmon, recreational activities, and commercial seafood processing. Potential for establishment was evaluated by examining water temperatures, spatial and temporal overlap of hosts, and the distribution and genetic composition of the oligochaete host, Tubifex tubifex. The most likely pathway of M. cerebralis transport in Alaska is human movement of fish by stocking. The extent of M. cerebralis infection in Alaskan salmonid populations is unknown, but if the parasite becomes dispersed, conditions are appropriate for establishment and propagation of the parasite life cycle in areas of south-central Alaska. The probability of further establishment is greatest in Ship Creek, where the abundance of susceptible T. tubifex, the presence of susceptible rainbow trout Oncorhynchus mykiss, and the proximity of this system to the known area of infection make conditions particularly suitable for spread of the parasite.     

Proteolytic activity in Hysterothylacium aduncum (Nematoda: Anisakidae), a fish gastrointestinal parasite of worldwide distribution

Proteases have a significant role in the life cycle of parasites and the pathogen-host relationship, being regarded as important virulence factors. In the parasitic nematode Hysterothylacium aduncum proteolytic activity was measured during in vitro development from third larval stage (L3) to mature adult, using DQ red casein as a fluorogenic substrate. Proteolytic activity was detected in all the developmental stages studied and at all pH values within the range employed (2.0-7.5). The assay with specific inhibitors permitted the determination of metalloprotease activity, and, to a lesser extent, that of aspartate- and cysteine-protease. Serine-protease activity was the lowest of those studied. In L3 recently collected from the host fish (L3-0 h), the greatest activity was found at an optimum pH of 4.0 and was mainly inhibited by 1,10-phenathroline (metalloprotease inhibitor). This metalloprotease activity in L3-0 h (infective stage) may be related to the invasion of the host tissues by this larva. In the other developmental stages, the greatest protease activity was found at pH 5.5, although at pH 4.0 a lower activity peak was detected. On the other hand, our data show that the proteolytic activity of the nematode varies according to the presence of pepsin (an aspartic-protease) in the culture medium. Thus, at pH 4.0, activity was greater in the absence of pepsin, with increasing aspartic-protease activity. Together with the detection of aspartic-, cysteine- and metallo-protease (enzymes involved in digestion in invertebrates) in all the developmental stages of the parasite taking place in the digestive tract of the host fish, this allows us to suggest that the pepsin in the culture medium mimics the predigestion conditions in the habitat of the worm within the host and that the activity detected may have, amongst others, a digestive function.     

Recent advances in the epidemiology,clinical and diagnostic features,and control of canine cardio-pulmonary angiostrongylosis

The aim of this review is to provide a comprehensive update on the biology, epidemiology, clinical features, diagnosis, treatment, and prevention of canine cardio-pulmonary angiostrongylosis. This cardiopulmonary disease is caused by infection by the metastrongyloid nematode Angiostrongylus vasorum. The parasite has an indirect life cycle that involves at least two different hosts, gastropod molluscs (intermediate host) and canids (definitive host). A. vasorum represents a common and serious problem for dogs in areas of endemicity, and because of the expansion of its geographical boundaries to many areas where it was absent or uncommon; its global burden is escalating. A. vasorum infection in dogs can result in serious disorders with potentially fatal consequences. Diagnosis in the live patient depends on faecal analysis, PCR or blood testing for parasite antigens or anti-parasite antibodies. Identification of parasites in fluids and tissues is rarely possible except post mortem, while diagnostic imaging and clinical examinations do not lead to a definitive diagnosis. Treatment normally requires the administration of anthelmintic drugs, and sometimes supportive therapy for complications resulting from infection.