Fostering in sheep. IV. Use of restraint

Fostering an additional lamb on to a ewe with a single lamb of her own was attempted in Dorset, Corriedale and Merino ewes restrained by the neck in pens for up to 12 days. Fostering occurred more rapidly in Dorsets than in Merinos, and success rates tended to be higher in Dorsets (81%) than the other breeds (73% in Corriedales and 69% in Merinos). The use of screens at the level of the ewe's neck to limit her olfactory examination of the lambs was more successful than when full olfactory contact was allowed. The prevention of any close olfactory contact resulted in an intermediate rate of acceptance, even though acceptance was apparently based on olfaction, as shown by universal rejection of strange alien lambs. Acceptance tests done within a few minutes of release from restraint were found not to reflect accurately the final level of acceptance, since some lambs came to be accepted within a day of being released. Further acceptances, that developed several days after release into a flock of ewes and lambs, emphasize an important role of the lamb in maintaining contact with the ewe. With few exceptions, lambs accepted when released from the pens were still accepted after a week or so in the flock.Dorset ewes given one or two foster lambs in the absence of their own lamb accepted the fosters within 9 days.Restraint fostering had no apparent harmful effects on the ewes or lambs, but had the disadvantages of being laborious and of soiling the ewes' wool. It has the advantage of providing lambs with an assured milk supply, and could prove useful for ewes that have failed to accept lambs under other methods.     

Fostering in sheep. I. Facilitation by use of textile lamb coats

Transfer of odour between lambs less than 1 week old was attempted by the use of rectangular coats of hessian (a jute textile) worn for 24 h by a ewe's own lamb and then transferred to a potential foster lamb. In the absence of their own lamb, 50–60% of ewes accepted the foster lamb immediately and more than 90% accepted the foster after confinement together over the next 2 days. Without exchange of coats, 12% of ewes accepted alien lambs immediately, and 50% after confinement.Breed differences in the success of fostering were observed, and a higher proportion of primiparas than multiparas accepted foster lambs. Foster lambs of similar appearance to the ewe's own lamb were more readily accepted than lambs of different appearance, and lambs 1–2 days old were more readily accepted than older lambs. There was a low level of success with fostering several weeks after lambing. Ewes that had accepted a foster lamb and then had their own lamb returned within 24 h of fostering initially tended to favour their own lamb.The success of the fostering method used in these studies appears to depend on both transfer and masking of lamb odour. The method should prove useful as an experimental tool, and as a practical means of saving lambs.     

Role of the vomeronasal organ in neonatal offspring recognition in sheep

Twenty-five pregnant Dorsett ewes were randomly divided into three groups to test if ewes use their vomeronasal organs for offspring recognition during nursing. One group of eight ewes (procaine) were made anosmic by irrigation of the nasal olfactory apparatus with a zinc sulphate procaine solution. The second group of nine ewes (cauterized) had their vomeronasal organs rendered nonfunctional by cauterization of the nasoincisive duct. The third group of eight ewes were the controls. Parturition was synchronized in all ewes with betamethasone on Day 145 of gestation. Maternal responsiveness was tested two separate times with 1- to 2-day-old alien lambs. Each alien lamb trial was conducted 24 h apart. Cauterized ewes allowed alien lambs to suckle and they were unable to distinguish alien lambs from their own lambs, whereas the ewes in both groups with functional vomeronasal organs (procaine and control) violently rejected any alien lamb's attempt to suckle. Thus, female sheep use their vomeronasal organs for neonatal offspring recognition.     

Mother–young bonding in prolific D’man and Romanov sheep

The effect of prolificacy on mother–young mutual recognition is still largely unknown in sheep. The aim of the present study was to investigate the ability of prolific ewes to develop an exclusive bond with their neonate and to recognize each other from alien subjects. Observations were performed on 11 D’man and 16 Romanov ewes with, respectively, 19 and 37 of their lambs. Maternal selectivity was tested at 24 h postpartum during two consecutive periods of 3 min. All mothers accepted their own lambs at suckling, while 21 out of 27 (78%) rejected the alien, thus displaying mostly selective nursing. To assess mother–young recognition, a two-choice test was performed at 36 h postpartum. Ewes clearly preferred their own lambs to an alien lamb and behavioral differences were not found between mothers of small (singletons and twins) and large litters (triplets and quadruplets). Romanov ewes reached their own lambsmore rapidly and spent more time near them than D’man ewes. Lambs from small litters, in particular light lambs, clearly preferred their mothers to an alien dam; however, both light and heavy lambs in large litters did not discriminate between the two stimulus ewes. D’man lambs clearly preferred their mothers to an alien dam; in contrast, such a preference was not as clearly demonstrated in Romanov lambs.     

Mother-lamb acoustic recognition in sheep: a frequency coding

Ewes of the domestic sheep ( Ovis aries ) display selective maternal investment by restricting care to their own offspring and rejecting alien young. This trait relies on individual recognition processes between ewes and lambs. Whereas identification at the udder is only olfactory, distance recognition is performed through visual and acoustic cues. We studied the effectiveness and modalities of mutual acoustic recognition between ewes and lambs by spectrographic analysis of their vocal signatures and by playbacks of modified calls in the field. Our results show that ewes and their lambs can recognize each other based solely on their calls. The coding of identity within the vocal signatures, previously unknown in sheep, is similar in lamb and ewe: it uses the mean frequency and the spectral energy distribution of the call, namely the timbre of the call. These results point out a simple signature system in sheep that uses only the frequency domain. This engenders a signal with low information content, as opposed to some highly social birds and mammal species that may integrate information both in the temporal and spectral domains. The simplicity of this system is linked to the roles played by vision and olfaction that corroborate the information brought by the vocal signature.     

Fostering in sheep. III. Facilitation by the use of odorants

Lambs were thoroughly anointed with various odoriferous substances to test the facilitatory effects on substitutive fostering on to multiparous Merino ewes held in individual pens. The ewes were first familiarised with an odorant by anointing their own lamb with it 24 h before a similarly anointed foster lamb was presented. The odorants were neatsfoot oil, wool wax, vegetable oil, butyric and propionic acids, mercaptoethanol, vanillin, methyl salicylate and eucalyptus oil. Acceptance of foster lambs was assessed over the next 48 h. Odorants were not applied to the ewes.All eight foster lambs treated with neatsfoot oil were accepted within 24 h; seven immediately. There were few immediate acceptances with the other odorants, but from two to five lambs out of groups of six were permanently fostered by 48 h. Only one foster lamb was accepted in the control group of 12 in which no odorant was used. Fostering was most successful with neatsfoot oil and wool wax, both substances that originated from ruminant skin.Fostering was less successful in tests that omitted pre-exposure of the ewes to their own lambs treated with neatsfoot oil, wool wax or vegetable oil.Toxicity problems were encountered with ethanol solutions of mercaptoethanol, methyl salicylate, eucalyptus oil and neatsfoot oil.     

Distance recognition in ewes and lambs kept permanently indoors or at pasture

We tested the ability of Merino and Merino × Romanov ewes and lambs to recognise each other at a distance when given the choice between an alien partner and their own. Two groups of animals were studied. In the first group, multiparous ewes and their lambs were housed permanently indoors at a stocking-rate of 1 ewe/2 m². In the second group, ewes and lambs were run permanently outside in a paddock at a stocking rate of 1 ewe/500 m². Of animals living outside, 87% of ewes recognised their own lamb (100% of multiparous dams versus 84% of primiparous ones, P = 0.02). This differed significantly from a random choice. By contrast, the proportion of correct choices by ewes living inside (all multiparous) was significantly lower (64%) and did not differ from a random choice at the 5% level. Lambs reared outside recognised their dam in 95% of cases, versus 72% in lambs reared indoors (P = 0.01). Both frequencies differed significantly from a choice at random. It is concluded that the environmental conditions in which ewes and lambs are maintained can influence their ability to select their partners at a distance with the help of auditory and visual cues.     

Pinealectomy delays puberty in ewe lambs

Fifteen pinealectomized and 15 unoperated ewes were exposed to constant light for 3 weeks before and 10 weeks after lambing. Fourteen pinealectomized and 15 unoperated ewes were allowed to lamb outdoors. Five ewe lambs born in constant light to the 2 groups of dams were pinealectomized at 10 weeks of age. Ewes and lambs were then returned to the field. Puberty (determined by weekly progesterone analysis) was significantly delayed (P less than 0.05) in the pinealectomized ewe lambs. Median pubertal age in pineal-intact ewe lambs was 37 weeks compared to 49 weeks in pinealectomized lambs. Constant light during the first 10 weeks of life had no effect upon puberty onset nor did the pineal status of the dam. Control lambs entered seasonal anoestrus at the time pinealectomized ewe lambs were entering puberty. Pinealectomized lambs entered anoestrus at the same time as control lambs were beginning their second breeding season. These results confirm a key role of pineal-mediated hormonal signals in the control of puberty in the sheep.     

Behavioural development in the neonatal lamb: effect of maternal and birth-related factors

The survival of the neonate relies on the integrated expression of appropriate behaviours from both the mother and young. In precocious species, like the sheep, the speed with which the lamb stands and seeks the udder is related to lamb survival. In this study the effects of birth difficulty, lamb birth weight, ewe loss or gain in backfat from conception to delivery, ewe parity, litter size, and lamb sex on neonatal lamb behavioural development were investigated in over 500 lambs of two breeds (Scottish Blackface and Suffolk). Lambs that required assistance to be delivered were significantly slower to perform all neonatal behaviours than unassisted lambs (P<0.001), and were less active over the first 3 days after delivery (P<0.05). There were no effects of lamb birthweight that were not accounted for by the increased likelihood of requiring assistance in heavier birth weight lambs. Ewes that mobilised less body fat during their pregnancy produced lambs that stood and sucked quickly (P<0.001), and were more active over the first 3 days of life. Lambs born to first parity ewes were slower to stand and suck than lambs born to experienced ewes (P<0.001). There was an improvement in time taken by lambs to stand, seek the udder and to suck with each increase in ewe parity. Litter size had an additional retarding influence on the behaviour of multiple-born lambs that could not be accounted for by birthweight. In the Suffolk breed male lambs were slower to stand and suck than female lambs, this effect was not seen in Blackface lambs. These data demonstrate that lambs that require assistance at birth, even if they survive the birth process, lambs born to ewes that lose a lot of condition over pregnancy or first parity ewes, triplet lambs and, at least in some breeds, male lambs are slower to progress through the sequence of neonatal behaviours. These lambs are, therefore, at greater risk of not surviving to weaning.     

Hearing recognition of the lamb by its mother

A group of twenty-three Merino ewes were tested for their ability to discriminate between their own and an alien lamb at a distance. Most of them (nineteen of twenty-three) were able to recognize their own young at a distance of 10 m. Eighteen of the twenty-three mothers, selected at random, were tested with recorded bleatings of lambs. They displayed a significant preference for the bleating of their own lambs. These results suggest that hearing plays an important role in the distance recognition of the lamb by its mother.     

Conditioning ewes and lambs to increase consumption of spotted knapweed

Spotted knapweed (Centaurea maculosa Lam.) is an invasive plant that alters species composition and grazing value of rangelands in the northwestern United States. The spread of invasive plants may be reduced by using livestock as a biological control. We determined if mature ewes and their lambs (n = 34 ewe/lamb pairs) consume more spotted knapweed when ewes and/or lambs are conditioned to fresh-cut spotted knapweed. Ewe/lamb pairs were randomly assigned to one of four conditioning treatments: ewes and lambs not conditioned to spotted knapweed (N), conditioned ewes with non-conditioned lambs (E), non-conditioned ewes with conditioned lambs (L), or conditioned ewes and lambs (both—B). Then, ewes and lambs were observed together for 5 days (Trial 1); 11 days later, lambs were observed for 4 days without their mothers (Trial 2). During conditioning, intake by conditioned and non-conditioned ewes and lambs varied over time (as-fed basis, treatment by day interaction; ewes P = 0.03; lambs P = 0.05). Overall, non-conditioned lambs (N, E) consumed more than conditioned lambs (L, B; P = 0.02). In Trial 1, N ewes consumed similar amounts of spotted knapweed and bromegrass (Bromus inermis Leyss.) as the E, L, and B ewes (P = 0.67). E ewes spent more time eating spotted knapweed than L ewes (P = 0.001), and E ewe/lamb pairs consumed more spotted knapweed than L ewe/lamb pairs (P = 0.02). In Trial 2, N lambs consumed less spotted knapweed than E, L, and B lambs (P = 0.06). L lambs consumed more than E lambs (P = 0.007). Conditioning ewes, lambs, or ewes and lambs did not increase time spent eating spotted knapweed when both grazed together in a drylot, but conditioned lambs, without their mothers present, consumed more spotted knapweed 11 days later than non-conditioned lambs. Conditioning lambs only in a group setting with their peers may have the greatest potential to enhance consumption of spotted knapweed, because of social facilitation and the predilection for young animals to try novel feeds.     

Congenital infection with Schistosoma japonicum but not with Schistosoma bovis in sheep

The present study investigated whether Schistosoma japonicum or Schistosoma bois could establish prenatally in lambs. Three ewes were exposed to S. japonicum by intramuscular injection of cercariae, and 3 ewes were exposed to S. bovis cercariae using the leg-emerging technique approximately 2 mo before delivery, and 1 age-matched pregnant ewe served as an uninfected control. The study lasted 18-20 wk after infection, which was 8-9 wk after delivery. All 6 exposed ewes became infected with either S. bovis or S. japonicum. Eight lambs were borne by the 7 ewes, of which 1 (S. bovis exposed) was dead and 1 (S. japonicum exposed) died at delivery. Of the 3 S. japonicum-exposed lambs, 2 were found infected. Four lambs born of S. bovis-exposed ewes were negative. Despite having no worms, these 4 S. bovis-exposed lambs as well as the 1 negative S. japonicum-exposed lamb had, in contrast to the nonexposed control lamb, few, but distinct, liver granulomas dominated by eosinophils and giant cells with large central necrotic areas but with no remnants of eggs or worms. Hence, congenital infection was demonstrated in S. japonicum-infected lambs, but not in S. bovis-infected ones.     

Effects of sex, litter size and periconceptional ewe nutrition on the ewe–lamb bond

Undernutrition restricted to the time around conception in sheep alters endocrine and metabolic responses in the offspring. Studies in rats suggest that such an early insult can also alter the behaviour of the offspring. We studied the effects of mild maternal periconceptional undernutrition (10–15% body weight reduction) on the lamb's response to separation from and reunion with the mother, and on the ewe–lamb bond, evaluated as the preference for each other over an alien ewe/lamb in a test enclosure, at 24 h, 1 and 4 weeks of age. Lamb birth weight was not affected by maternal nutrition. Maternal periconceptional undernutrition did not affect the lambs’ responses to separation and reunion (number of vocalisations, times to leave pen and achieve proximity with ewes) or the bond between ewes and lambs (percentage of time spent near their own dam/lamb). However, there were effects of sex, litter size and time on lambs’ responses to separation and reunion and on the ewe–lamb bond. Female lambs vocalised more during separation (P ≤ 0.01) and left the holding pen faster than males (P ≤ 0.05), and twins vocalised more than singletons (P ≤ 0.05). The time to leave the holding pen, achieve proximity with the ewes and number of vocalisations decreased with increasing postnatal age of the lamb (all P ≤ 0.001). Ewes showed a stronger preference for the lamb at 24 h and 1 week than at 4 weeks postpartum (P ≤ 0.001), whereas lambs followed an opposite trend, showing a lower preference for the mother at 24 h and 1 week and increased preference at 4 weeks of age (P ≤ 0.001). We conclude that periconceptional undernutrition does not affect offspring response to separation from the mother or the bond between ewes and lambs, but that these behaviours are affected by sex and litter size for up to 4 weeks after birth.     

The effects of ablation of the olfactory bulbs on parturition and maternal behaviour in Soay sheep

Eight pregnant Soay ewes were made anosmic by ablation of the olfactory bulbs before their lambs were born. They were observed during parturition and in the following few days. Four out of five ewes seen during parturition did not show pre-natal lip licking which is normally seen in Soays. Four ewes did not lick their lambs properly after birth, and in six there was a lack of specific response to their own lambs, so that strange lambs were often fed. Post-mortem examination showed that olfactory tracts were absent in all except one of the ewes examined. It is suggested that the importance of olfaction in maternal care is concerned with identification of the lamb at close quarters and in forming the ewe-lamb relationship.     

Selection for reproductive rate in Rambouillet sheep: Estimated genetic change in reproductive rate

Rambouillet sheep were selected for high or low reproductive rate based on an index of reproductive rate, reproductive INDEX = dam's total lifetime lambs born/(age in years-1). Selected ewes were first born in 1969. A random bred control line was established from the remaining foundation ewes with the first ewes born in 1973. Genetic variances were estimated with REML procedures for reproductive index and number of lambs born or weaned either per ewe exposed for breeding or per ewe lambing. Heritability of reproductive index was 0.26. Heritability for number of lambs born were 0.11 and 0.12 and for number of lambs weaned were 0.04 and 0.04 per ewe exposed for breeding and per ewe lambing, respectively. Breeding values for the reproductive index were estimated using the full animal model (BLUP) with the complete numerator relationship matrix on reproductive index, with fixed effect of year of birth of ewe and breeding values for number of lambs born or weaned either per ewe exposed for breeding or per ewe lambing, estimated with the fixed effects of year of birth of ewe and age of ewe and the random effect of permanent environmental effect among repeated lambing records. Genetic change was estimated as the regression of breeding value for reproductive index and number of lambs born or weaned either per ewe exposed for breeding or per ewe lambing on year of birth of the ewe. High and low lines both responded to selection for reproductive index as compared to the control line (P<0.01). The high line increased at a rate of b = 0.0134 ± 0.0006 reproductive index units and the low line decreased at a rate of b = − 0.0098 ± 0.0005 reproductive index units per year. Response to selection for reproductive index resulted in b = − 0.0074 ± 0.0007 and b = 0.0163 ± 0.0006 lambs per ewe exposed for breeding, and b = − 0.0041 ± 0.0002 and b = 0.0075 ± 0.0002 lambs weaned per ewe exposed for breeding in the low and high lines, respectively. There was a greater response to selection in the high line compared to the low line. These results indicated that the reproductive index did respond to selection.     

Amniotic fluid is important for the maintenance of maternal responsiveness and the establishment of maternal selectivity in sheep

Amniotic fluid (AF) is important for the establishment of maternal behaviour in inexperienced ewes, but its role in experienced mothers remains to be studied. Here, the maintenance of post-partum maternal responsiveness and the establishment of exclusive bonding was investigated in multiparous ewes when AF was removed from the neonate or/and physical contact with the young was precluded for the first 4 h post partum. Maintenance of maternal responsiveness and establishment of exclusive bonding were measured by the proportion of mothers accepting their own lamb and alien lambs that had been either washed or not washed, and by comparing an acceptance score for each type of lamb. The acceptance score was computed by summing standardised variables of acceptance (low bleats, acceptance at udder, nursing and licking time) and subtracting standardised variables of rejection (high-pitched bleats, rejection at the udder and aggressive behaviour). Washing the neonate reduced its acceptance score, but the proportion of mothers rejecting their own lamb was reduced only when washing the neonate and prevention of physical contact for 4 h were combined (7/15 v. 0/10 in controls, P = 0.02). In addition, washing the neonate increased the acceptance score of the washed alien lamb, but not of the unwashed alien. However, washing and privation of physical contact did not increase significantly the proportion of mothers accepting an alien lamb at 4 h post partum. We conclude that AF is important in experienced ewes for the establishment of maternal responsiveness, as already found in primiparous mothers. In addition, our results indicate that AF also carries some chemosensory information facilitating exclusive bonding.     

Comparison of the survival of fertilized eggs from adult ewes in the uteri of adult ewes and ewe lambs

Fertilized eggs, obtained from mature donor ewes, were transferred into the uteri of adult ewes and ewe lambs (1 or 2 eggs per recipient). The survival rate to term of the transferred eggs was similar in the two classes of recipients. The percentage of adult ewe and ewe lamb recipients which gave birth to twins was 45.3 and 41.1 respectively. Gestation length was shorter (P less than 0.01) and lamb birth weight lower (P less than 0.01) for ewe lamb recipients. The results indicate that the generally lower lambing rate of ewe lambs compared to adults is unlikely to be due to unfavourable uterine conditions.     

Maternal selectivity suppression through peripheral anosmia affects neither overall nursing frequency and duration, nor lactation performance in ewes

The effects of prepartum peripheral anosmia on nursing activity, milk production and growth of the lambs, were assessed by comparing intact (n=10) and anosmic (n=10) multiparous Columbia and Rambouillet ewes and their single lamb during the first 2 months of lactation. Intact mothers only nursed their own lamb (98%) while most of the nursing activity in anosmic mothers concerned alien lambs (78%). On the other hand, the total duration and the frequencies of nursing did not differ significantly between groups (P>0.05). Nevertheless, the total percentage of nursing of own lamb by anosmic mothers (22%) was higher than expected at random (10%). Milk production or lambs' weights did not differ between groups. We conclude that prepartum anosmia resulted in the failure of ewes to develop true selective nursing up to the 8th week of lactation, although some preferential mother-young relationship yet developed. On the other hand, it did not affect significantly overall nursing activity.     

Effect of plane of nutrition of 1- and 2-year-old ewes in early and mid-pregnancy on ewe reproduction and offspring performance up to weaning

The objective of the present study was to determine the effect of plane of nutrition in early pregnancy (EP) and mid-pregnancy (MP), on the productive performance of 1- and 2-year-old ewes and their offspring. Over 2 successive years, between days 0 and 39 after synchronized mating (EP), 1- (n = 117) and 2- (n = 52) year-old ewes were allowed 60% (low, L-EP), 100% (medium, M-EP) or 200% (high, H-EP) of requirements for maintenance (M). Between days 40 and 90 (MP), 1-year-old ewes were allowed 140% (M-MP) or 200% (H-MP), while 2-year-old ewes were allowed 80% (M-MP) or 140% (H-MP) of their M requirement. After day 90, all ewes were fed to meet requirements for late pregnancy. Increasing the plane of nutrition between days 0 and 39 resulted in increases in live weight (LW) (P < 0.001) and body condition score (BCS) (P < 0.001) during the EP period (H-EP > M-EP > L-EP), differences that in 1-year-old ewes were sustained to lambing (P < 0.05). On day 42 of gestation H-EP ewes had lower plasma progesterone concentrations than L-EP or M-EP ewes in 1- (P < 0.01) and 2- (P < 0.001) year olds. This was concomitant with diet H-EP tending to reduce the number of lambs born per ewe in both age groups (P = 0.06 and 0.07, respectively). Foetuses from 1-year-old L-EP ewes had smaller cranial (P < 0.01) and abdominal (P < 0.05) diameters at day 53 of gestation, with H-EP lambs tending to be heaviest at birth (P = 0.07). Similar findings were recorded for 2-year-old ewes. One-year-old ewes offered diet L-EP presented negative maternal behaviours more frequently (P < 0.05), while the incidence of lamb mortality at 6 weeks tended to be greater for L-EP lambs (P = 0.07). In MP, 1-year-old ewes offered diet M-MP were associated with foetuses with bigger abdominal diameters at day 78 (P < 0.05). However, there were no differences in lamb weight or size at term (P > 0.05). These ewes exhibited more positive maternal behaviours (e.g. increased grooming frequency and duration; P < 0.05) than ewes offered diet H-MP, and their offspring were more successful in suckling (P < 0.05). Results suggest that in young ewes, a temporary nutrient restriction in EP resulted in increased prolificacy. However, ewes and their offspring were lighter at birth and ewe maternal behaviour was poorer, resulting in increased lamb mortality. In MP, a medium plane of nutrition offered to 1-year-old ewes led to improved maternal and offspring behaviour.     

Gastro-intestinal nematode parasite populations in ewes and lambs and the origin and time course of infective larval availability in pastures

Changes in the numbers of infective larvae available on pasture and the development of helminth infection in spring-born lambs were studied in an environment with a mild winter and where pastures are grazed throughout the year. Prior contamination of pastures in autumn and early winter was compared with egg deposition by lambing ewes as sources of infection. Larvae derived from eggs deposited by lambing ewes did not appear on pasture until 8 weeks after the beginning of lambing but were present in increasing numbers for the last 7 weeks before weaning at 11–15 weeks of age. Lambs exposed to prior contamination became infected earlier than those exposed only to ewe contamination, but lamb egg output made no contribution to larval numbers on pasture before weaning. Lamb growth rates to weaning did not differ between groups exposed either to prior contamination alone, to ewe egg output alone or to both, but all groups carried appreciable infections at weaning. After weaning, lambs remained on the same pastures, and those exposed to ewe contamination suffered severe trichostrongylosis 5 weeks after weaning. Lambs exposed only to over-wintered contamination also developed severe trichostrongylosis 9 weeks after weaning. The results are compared with those of similar recent studies in Britain and their implications for control of infection are discussed.