Responses of hair and wool sheep to a single fixed dose of infective larvae of Haemonchus contortus

Body weights, fecal egg counts (FEC), and packed cell volumes (PCV) of hair sheep and wool sheep crossbred lambs were compared over 8 weeks following administration of a single dose of approximately 10,000 third-stage larvae of Haemonchus contortus. Hair sheep lambs (n=17) were reciprocal crosses between mainland USA populations of Barbados Blackbelly (BB) and Virgin Islands White (VIW) sheep. Wool sheep lambs (n=64) were from a crossbred composite of 50% Dorset, 25% Rambouillet, and 25% Finnish Landrace breeding. Lambs of both breed types continued to grow during the period of infection. Mean weights were higher for wool lambs (39.7±0.8 kg) than for hair lambs (28.2±1.5 kg). FEC increased to week 5 in both breed groups and remained elevated in wool lambs through week 7 but declined sharply in hair lambs after week 5. Mean FEC for weeks 4 through 8 were 4011±361 eggs per gram of feces (epg) in wool lambs, and 1135±196 epg in hair lambs. PCV declined through week 7 in wool lambs but stabilized and then increased after week 4 in hair lambs. Mean PCV in weeks 4 through 8 were 22.4±0.3% in wool lambs and 24.3±0.5% in hair lambs. These results suggest that Caribbean hair breeds may be able to contribute significantly to development of parasite-resistant sheep populations.     

Reciprocal Allogrooming in Wild Impala Lambs

Adult impala engage in a form of reciprocal allogrooming distinguished by a high degree of reciprocity and ***lack of influence of dominance or relatedness on partner preference or distribution of grooming between partners. A previous study on reciprocal allogrooming of captive newborn impala lambs in a zoological park found that the allogrooming emerged as early as the first week after birth and was identical in structure and reciprocity to allogrooming in adults. Because these findings of apparently unique allogrooming behavior of newborn impala could have been a reflection of the effects of being born and raised in a small, stable captive herd, it was necessary to investigate reciprocal allogrooming in newborn impala in the wild. The emergence, reciprocity, rate, and partner distribution of reciprocal allogrooming in wild newborn impala were observed at two study sites: a national park in Zimbabwe and a game farm in South Africa. Maternal one-way grooming between mother and newborn emerged as distinct from reciprocal allogrooming and rapidly declined after week 1 postpartum. Reciprocal allogrooming by lambs was first seen between 5 and 8 d postpartum, and as soon as the behavior occurred it was the same basic pattern as seen in adult impala. The reciprocity index for lambs was near 0.5, indicating that lambs delivered as much grooming during an encounter as the partner. Lambs were grooming frequently with non-mother adults and other lambs by week 1 or 2; by week 3 and onward the allogrooming rate of lambs was more than twice that of their mothers, as predicted by the body size principle of the programmed grooming model. The strong predisposition of neonatal impala lambs to deliver reciprocal allogrooming as early as the first or second week postpartum would appear to reflect a genetically acquired adaptation to the threat of tick infestation in their natural habitat.     

A functional analysis of social grooming patterns through direct comparison with self-grooming in rhesus monkeys

Social grooming in primates is a complex behavior in which monkeys stroke, pick, or otherwise manipulate a companion’s body surface. While grooming has been associated with important social functions, researchers who have examined its physical characteristics, such as body site preferences, have focused on its role in skin care and ectoparasite removal. Whether the form of social grooming is constrained primarily by utilitarian or social functions was empirically tested by directly comparing this behavior with self-grooming, which was assumed to have primarily utilitarian functions. Predictions made concerning social grooming, based on a comparison with self-grooming, were tested by examining site preferences, duration of grooming (overall and to specific body sites), and method of grooming (stroking, picking, or a combination of the two) in rhesus monkeys. None of these predictions was verified. The physical characteristics of social grooming could not be predicted from the way an animal groomed itself. There was no relationship between site preferences in these two types of grooming. Animals groomed others for longer periods of time than they groomed themselves, both overall and when grooming specific sites. The methods these animals used differed markedly in self-and social grooming: they primarily picked at their own hair or skin but, when grooming others, varied their technique across sites, stroking the hair where it was thickest and picking where the hair was sparse. These results suggest that utilitarian functions are not the most important factors constraining the form of social grooming.     

How Did Domestication Change the Hair Morphology in Sheep and Goats?

We analysed macro-and microscopic features of dorsal guard hairs in 21 specimens of wild and domestic sheep and goats. We integrated and extended the available data on hair morphology of wild species and provide a first comparative analysis of hair structure of domestic forms. Domestic sheep and goats, probably due to a convergence process under artificial selection, show similar medullary features to each other and different medullary structures from their relative wild relatives. Different breeds show a diverse alteration of the medullary structure probably correlated to the duration of the domestication process. Domestic sheep have a cuticular structure different from the relative wild ancestor, while domestic goats do not show clear differences in the cuticle from the related wild species. The strong artificial selection for wool production may have transformed the hair structure of the sheep, but not that of the goat. We described the effects of age on the microscopic structure of hair, which have not yet been investigated. The medullary structure and the cuticular pattern in domestic forms do not change with age, as seen in wild species, because juveniles characters are retained in adults due to domestication.     

New Mechanism of Melanosomes Transport into Hair of Lambs of Different Breeds and Genotypes

By light microscopic investigation of skin and wool specimens of newborn lambs, we discovered a previously unknown mechanism for melanosomes transport in the process of dermal papilla melanocytes regular mitosis and migration into the hair shaft. This mechanism plays a great role in hair pigmentation especially in dominant (E^D/E^D) and recessive (A^a/A^a) black lambs of all investigated breeds. The rate of pigment cell mitosis, proliferation, and migration differs greatly in lambs of investigated color genotypes. In black genotypes the rate of melanocyte mitosis is very high and is approximately the same as in the hair bulb matrix cells, whereas in brown and red genotypes this rate is much lower. Melanocyte mitosis in the light red and tan groups was not found.     

New mechanism of melanosomes transport into hair of lambs of different breeds and genotypes.

By light microscopic investigation of skin and wool specimens of newborn lambs, we discovered a previously unknown mechanism for melanosomes transport in the process of dermal papilla melanocytes regular mitosis and migration into the hair shaft. This mechanism plays a great role in hair pigmentation especially in dominant (ED/ED) and recessive (Aa/Aa) black lambs of all investigated breeds. The rate of pigment cell mitosis, proliferation, and migration differs greatly in lambs of investigated color genotypes. In black genotypes the rate of melanocyte mitosis is very high and is approximately the same as in the hair bulb matrix cells, whereas in brown and red genotypes this rate is much lower. Melanocyte mitosis in the light red and tan groups was not found.     

Exposure to ram wool stimulates gonadotropin-releasing hormone pulse generator activity in the female goat

In the sheep and goat, exposure of anestrous females to a conspecific male odor enhances reproductive activity. Interestingly, a previous report indicated that male goat hair stimulated pulsatile luteinizing hormone (LH) secretion in the ewe. In the present study, we addressed whether ram wool affects the gonadotropin-releasing hormone (GnRH) pulse generator activity in the female goat. Five ovariectomized (OVX) goats were chronically implanted with recording electrodes in the mediobasal hypothalamus, and manifestations of the GnRH pulse generator were monitored as characteristic increases in multiple-unit activity (MUA volleys). Wool or hair samples were collected from a mature ram, ewe and male goat, and their effects on the MUA volley were examined. The exposure to ram wool induced an MUA volley within 1 min in all five OVX goats, as did the exposure to male goat hair. The ewe wool had no effect on the timing of an MUA volley occurrence. An invariable association of MUA volleys with LH pulses in the peripheral circulation was also confirmed in two OVX goats exposed to ram wool. The present results clearly indicate that exposure to ram wool stimulates pulsatile GnRH/LH release in the female goat. Since exposure to male goat hair enhances pulsatile LH secretion in the ewe, it is likely that very similar, if not identical, molecules are contained in the male-effect pheromone in the sheep and goat.     

Regulation of brain water during acute hyperosmolality in ovine fetuses, lambs, and adults.

In adult rats, when plasma osmolality increases, water flows across the blood-brain barrier down its concentration gradient from brain to plasma, and brain volume deceases. The brain responds to this stress by gaining osmotically active solutes, which limit water loss. This phenomenon is termed brain volume (water) regulation. We tested the hypothesis that brain volume regulation is more effective in young lambs and adult sheep than in fetuses, premature lambs, and newborn lambs. Brain water responses to acute hyperosmolality were measured in the cerebral cortex, cerebellum, and medulla of fetuses at 60 and 90% of gestation, premature ventilated lambs at 90% of gestation, newborn lambs, young lambs at 20-30 days of age, and adult sheep. After exposure of the sheep to increases in systemic osmolality with mannitol plus NaCl, brain water content and electrolytes were quantified. The ideal osmometer is a system in which impermeable solutes do not enter or leave in response to an osmotic stress. There were significant differences from an ideal osmometer in the cerebral cortex of fetuses at 90% of gestation, cerebral cortex, and cerebellum of newborn lambs, and cerebral cortex, cerebellum, and medulla of young lambs and adult sheep; however, there were no differences in the brain regions of fetuses at 60% of gestation and premature lambs, cerebellum and medulla of fetuses at 90% of gestation, and medulla of newborn lambs. We conclude that 1) brain water loss is maximal and brain volume regulation impaired in most brain regions of fetuses at 60 and 90% of gestation and premature lambs; 2) brain volume regulation develops first in the cerebral cortex of the fetuses at 90% of gestation and in the cerebral cortex and cerebellum of newborn lambs, and then it develops in the medulla of the lambs at 20-30 days of age; 3) brain water loss is limited and volume regulation present in the brain regions of young lambs and adult sheep; and 4) the ability of the brain to exhibit volume regulation develops in a region- and age-related fashion.     

The social grooming of captive female rhesus monkeys: Effects of the births of their infants

We observed the grooming interactions of 13 female rhesus monkeys (Macaca mulatta)before and for 12 weeks after the births of their infants. Mothers groomed for similar amounts of time before and after the birth of their infants, but after the birth, the grooming they directed to their infants may have been at the expense of that directed to other partners. Lactating females did not receive more grooming from other females but were approached more often, suggesting that they were more attractive. Mothers that groomed their infants most groomed others least, as if grooming time was limited for each mother or as if she was trying to compensate for avoiding interactions with other partners. Mothers of male infants groomed others more than mothers with female infants did, which might be due to mothers with daughters receiving more aggression and therefore avoiding interaction. Experienced and high-ranking mothers groomed their newborn infants considerably more than primiparous mothers did in the 24 hr following birth. Grooming was preferentially directed at close kin before the births of the infants. Mothers tended to groom higher-ranked partners more than they were groomed by them, and they tended to receive more grooming from lower-ranked partners than they gave, as suggested in models of rank attractiveness.     

Hemoglobin function in the developing goat and sheep

Comparative studies of the blood of newly born goats and sheep have indicated a number of mechanisms which are responsible for a decreasing affinity for molecular oxygen in these developing animals during the first 40 to 60 days after birth.
The concentration of 2,3-DPG in the red cells of young goats increases four to sixfold during the first 3 to 4 days of life, and this increase is associated with a marked decrease in the cellular pH; 2,3-DPG does not bind to hemoglobins of goats and the decreased affinity for oxygen of goat blood at this period is apparently due to the lowered pH produced by the large increase of intracellular anions. Similar changes occur in young lambs.
After 15 to 20 days the changes of the dissociation curve are related more to structural differences between adult and fetal hemoglobins; cellular pH moves closer to the values of adult red cells. In goats of this age the predominate hemoglobins are those with β chains and these have dissociation curves shifted further to the right than other adult hemoglobins.
In young lambs Hb-C is found only in association with Hb-A but the amount present seldom exceeds 5 to 10%. The oxygen affinities of sheep Hb-A and Hb-C are identical but higher than that of sheep Hb-B.     

Grooming in desert bighorn sheep (Ovis canadensis mexicana) and the ghost of parasites past

Ectoparasites such as ticks have a negative effect on host fitness,whereas parasite-defense grooming is effective in removing ticks.The central control (programmed grooming) model proposes thatanimals engage in preventive tick-defense grooming in responseto an internal timing mechanism, even in the absence of peripheralstimulation from parasites. This model predicts that smalleranimals will groom more frequently than larger ones becauseof the higher cost of parasitism for a small animal (body sizeprinciple). The peripheral stimulation (stimulus driven) modelpredicts no size-related differences in grooming rate in theabsence of tick bite irritation. We observed grooming behaviorin a Chihuahuan desert population of bighorn sheep (Ovis canadensismexicana), where ticks have been absent for perhaps thousandsof years. Although not exposed to ticks, bighorns self groomedby means of oral and scratch grooming, albeit at very low ratescompared to size-matched ungulates in both tick-infested andtick-free environments. Logistic regression and general linearmodels revealed both the probability that grooming was performedduring a 10-min focal sample and the rate of grooming when itoccurred was greater for younger, smaller age/sex categoriesof less body mass. Oral and scratch grooming were negativelyassociated with body mass during both years, with juveniles(X = 15 kg) grooming the most frequently and the oldest males(X = 70–85 kg) grooming the least. Assuming that programmedgrooming evolved in a tick-infested environment, the currentgrooming behavior of this population is a relict of their ancestralenvironment, an adaptation to the "ghost of parasites past."     

Attempted Cross-Transmission of Coccidia Between Sheep and Goats and Description of Eimeria ovinoidalis sp. n.

SYNOPSIS.
Attempted infection of 2 young lambs with oocysts of Eimeria christenseni from a goat was unsuccessful. Negative results were obtained also when young kids were fed oocysts of Eimeria ninakohlyakimovae from sheep. There was no difficulty in infecting lambs with the sheep coccidium resembling E. ninakohlyakimovae nor goats with the goat coccidium E. christenseni. Oocysts from the goat measured 38.4 × 26.7 m, but were easily distinguished from Eimeria ahsata from the sheep by sporocyst size and shape, and from Eimeria ovina by oocyst size. Eimeria ninakohlyakimovae -like oocysts from sheep averaged 23.0 ×18.2 m and were morphologically indistinguishable from previously reported goat coccidia.
Since no cross infections of sheep and goats could be accomplished with oocysts of Eimeria sp. characteristic of one or the other host, I concluded that sheep coccidia previously known as E. ninakohlyakimovae are distinct from morphologically similar goat coccidia and therefore constitute a separate species. Since the name E. ninakohlyakimovae was first used for coccidia from the goat, the sheep coccidium is renamed Eimeria ovinoidalis with oocyst structure and endogenous stages similar to those previously described from the sheep.     

Peripolar cell hypertrophy in the renal juxtaglomerular region of newborn sheep

Summary In the renal juxtaglomerular region of newborn sheep, it was found that glomerular peripolar cells and their granules were very much larger than those found in fetal lambs or adult sheep. Similar peripolar cell hypertrophy was triggered in fetal lambs treated in utero with intraperitoneal injections of dexamethasone. Ultrastructurally, granules of peripolar cells from newborn lambs resembled closely the enlarged zymogen granules described in the pancreas of newborn rats. Such peripolar cell hypertrophy may reflect a functional adaptation of the kidney to immediate postnatal life.     

Sexually and Developmentally Dimorphic Grooming: A Comparative Survey of the Ungulata

Grooming is effective in removing fitness‐compromising ectoparasites, such as ticks. Tick‐removal grooming is regulated both by a central control mechanism (programmed grooming model) and by cutaneous stimulation from tick bites (stimulus‐driven model). The programmed grooming model predicts that organismic factors that impact the cost‐benefit ratio of grooming (e.g., sex and body size) will influence the rate of grooming. The ‘vigilance principle’ predicts that breeding males of sexually dimorphic species will groom less than females to maintain high levels of vigilance for rival males and oestrus females. The intraspecific body size principle predicts that juveniles will engage in more frequent grooming than larger‐bodied adults to compensate for the higher cost of tick infestation for smaller animals. To examine the generalizability of these predictions, we surveyed the grooming rate of 53 species/subspecies of ungulates (primarily Bovidae and Cervidae) in tick‐free zoological parks in which stimulus‐driven grooming was controlled for. Matched‐pair analysis supported both predictions. Males delivered fewer oral grooming episodes per hour than females in 85% of sexually dimorphic species (n = 40), but only 56% of monomorphic species (n = 11) exhibited sexually dimorphic grooming, which was not different from random. Juvenile oral episodes per hour exceeded that of adult females in 74% of surveyed species (n = 36). As predicted by the body size principle, the grooming rate of juveniles declined as juveniles grew larger and less vulnerable to tick infestation. In separate analyses of Bovidae and Cervidae to control for common ancestry, both families supported sexually dimorphic grooming, but developmentally dimorphic grooming was supported only for Bovidae. Our results indicate that sexually dimorphic grooming is widespread in the ungulate species surveyed, suggesting that programmed grooming evolved at least as early as the common ancestor to the Artiodactyla.     

Testing the interspecific body size principle in ungulates: the smaller they come, the harder they groom

Tick removal grooming may be centrally regulated by an internal timing mechanism operating to remove ticks before they attach and engorge (programmed grooming model) and/or evoked by cutaneous stimulation from tick bites (stimulus-driven model). The programmed grooming model predicts that organismic and environmental factors that impact the cost-benefit ratio of grooming (e.g. body size and habitat) will influence the rate of tick removal grooming. The body size principle predicts that smaller-sized animals, because of their greater surface-to-mass ratio, should engage in more frequent tick removal grooming than larger-bodied animals in order to compensate for higher costs of tick infestation. The body size principle may be tested intraspecifically between young and adult animals, or interspecifically among species of contrasting body sizes. To rigorously test the interspecific body size prediction, we observed the programmed grooming (oral and scratch grooming) of 25 species (or subspecies) of bovids at a tick-free zoological park in which stimulus-driven grooming was ruled out. Multiple correlation analysis revealed highly significant negative correlations between species-typical mass and mean species grooming rates when habitat was controlled for in the model. Species-typical habitat type (classified along a gradient from most open to most closed) was positively correlated with mean oral grooming rate, indicating that species tended to groom at a higher rate in woodland and forest habitats (where typical tick density would be high) compared with more open environments. Species mass accounted for up to two-thirds of the variation in grooming rate across species, whereas habitat accounted for ca. 20% of variation in oral grooming. Similar results were obtained when the analysis was expanded to include 36 species/subspecies of six different families. The body size principle can therefore account for a large proportion of species-typical differences in programmed grooming rate among ungulates. However, to understand the tick defence adaptations of very large mammals that rarely or never engage in oral or scratch grooming (e.g. elephants, giraffes, rhinoceros), alternative tick defence strategies must be considered, such as thick skin, wallowing, rubbing and tolerance of oxpeckers and other tick-eating birds. Copyright 2000 The Association for the Study of Animal Behaviour.     

The evolution of parasite-defence grooming in ungulates

Grooming repertoires are exhibited by all terrestrial mammals, and removal of ectoparasites is an important ancestral and current function. Parasite‐defence grooming is regulated both by a central control mechanism (programmed grooming model) and by cutaneous stimulation from bites (stimulus‐driven model). To study the evolution of parasite‐defence grooming in ungulates, we compared species‐typical grooming behaviour with host morphology and habitat to test predictions of the programmed grooming model while taking into account phylogenetic relatedness. We observed grooming in 60 ungulate species at ectoparasite‐free zoological parks in which the confound of differential tick exposure was controlled for and stimulus‐driven grooming was ruled out. Concentrated‐changes tests indicated that sexually dimorphic grooming (in which breeding males groom less than females) has coevolved with sexual body size dimorphism, suggesting that intrasexual selection has favoured reduced grooming that enhances vigilance of males for oestrous females and rival males. Concentrated‐changes tests also revealed that the evolution of complex oral grooming (involving alternate use of both teeth and tongue) and adult allogrooming (whereby conspecifics oral groom body regions not accessible by self grooming) was concentrated in lineages inhabiting closed woodland or forest habitat associated with increased tick exposure, with such advanced grooming being most concentrated in Cervidae. Regression analyses of independent contrasts indicated that both host body size and habitat play a role in the evolution of species‐typical oral grooming rates, as previously reported. These results indicate that the observed grooming represents centrally driven grooming patterns favoured by natural selection in each lineage. This is the first phylogenetically controlled comparative study to report the evolution of parasite‐defence grooming behaviours in response to selection pressures predicted by the programmed grooming hypothesis. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 17–37.     

Grooming as a reward? Social function of grooming between females in cooperatively breeding marmosets

Classical models of grooming predict that subordinate primates will direct grooming towards dominants to receive coalitionary support from them. In contrast, recent reviews suggest that grooming asymmetries can change with social system and ecological conditions and should reflect asymmetries in services provided by different members of the dyad. We studied grooming patterns between females in six wild groups of common marmosets, Callithrix jacchus, to investigate the relation between social structure and grooming between females in a cooperatively breeding species. We observed grooming frequently and consistently in all study groups. Breeding females groomed nonbreeding females more than vice versa, and grooming between breeding and nonbreeding females was not related to agonistic behaviour. Our results provide some support to the hypothesis that grooming asymmetries are related to differences in services provided by different group members. We suggest that, in cooperatively breeding systems, breeding females may use grooming as an incentive for helper females to stay in the group.     

Comparison of cleft palate induction by Nicotiana glauca in goats and sheep

The induction of cleft palate by Nicotiana glauca (wild tree tobacco) during the first trimester of pregnancy was compared between Spanish-type goats and crossbred western-type sheep. Cleft palate was induced in 100% of the embryonic/fetal goats when their pregnant mothers were gavaged with N. glauca plant material or with anabasine-rich extracts from the latter, during gestation days 32-41. Seventy-five percent of newborn goats had cleft palate after maternal dosing with N. glauca during gestation days 35-41, while no cleft palates were induced when dosing periods included days 36-40, 37-39, or day 38 only. The induced cleft palates were bilateral, involving the entire secondary palates with complete detachment of the vomer. Eleven percent of the newborn goats from does gavaged during gestation days 32-41 had extracranial abnormalities, most often contractures of the metacarpal joints. Most of these contractures resolved spontaneously by 4-6 weeks postpartum. One newborn kid also had an asymmetric skull due to apparent fetal positioning. No cleft palates were induced in lambs whose mothers were gavaged with N. glauca plant or anabasine-rich extracts during gestation days 34-41, 35-40, 35-41, 36-41, 35-51, or 37-50. Only one of five lambs born to three ewes gavaged with N. glauca plant material during gestation days 34-55 had a cleft palate, but all five of these lambs had moderate to severe contractures in the metacarpal joints. The slight to moderate contracture defects resolved spontaneously by 4-6 weeks postpartum, but the severe contractures resolved only partially. Embryonic/fetal death and resorption (determined by ultrasound) occurred in 25% of pregnant goats fed N. glauca compared to only 4% of pregnant sheep. Nicotiana glauca plant material contained the teratogenic alkaloid anabasine at 0.175% to 0.23%, dry weight, demonstrating that Spanish-type goats are susceptible to cleft palate induction by the natural toxin anabasine, while crossbred western-type sheep are resistant. However, clinical signs of toxicity were equally severe in goats and sheep, even though maternal alkaloid tolerance was generally lower in sheep. We postulate that an alkaloid-induced reduction in fetal movement during the period of normal palate closure is the cause of the cleft palate and multiple flexion contractures. Teratology 61:203-210, 2000. Published 2000 Wiley-Liss, Inc.     

Comparison of sandbathing and grooming in two species of kangaroo rat

Sandbathing and grooming behaviours of two species of kangaroo rat (Dipodomys merriami and D. microps) were compared after zero and three days of sand deprivation. Results showed that D. merriami sandbathed and groomed at higher frequencies and over a longer time period than D. microps. Comparison of hair lipids revealed that D. merriami produced more oils. The data support the hypothesis that lipids in the hair are removed by sandbathing and suggest that D. merriami requires more pelage maintenance than D. microps. Three hypotheses about the function of lipids in the pelage of kangaroo rats are discussed.     

The Chimpanzee''s service economy: Food for grooming

Evidence is presented that the reciprocal exchange of social services among chimpanzees (Pan troglodytes) rests on cognitive abilities that allow current behavior to be contingent upon a history of interaction. Food sharing within a captive colony of chimpanzees was studied by means of 200 food trials, conducted on separate daus over a 3-year period, in which 6,972 approaches occurred among the nine adults in the colony. The success rate of each adult, A, to obtain food from another adult, B, was compared with grooming interactions between A and B in the 2 hours prior to each food trial. The tendency of B to share with A was higher if A had groomed B than if A had not done so. The exchange was partner-specific, i.e., the effect of previous grooming on the behavior of food possessors was limited to the grooming partner. Grooming did not affect subsequent sharing by the groomer, only by the groomee. The effect of grooming was greatest for pairs of adults who rarely groomed. Nevertheless, the effect was general: 31 dyadic directions showed an increase in sharing following grooming, and only 11 a decrease. Food possessors actively resisted approaches by individuals who had not groomed them. After food trials there was a significant reduction of grooming by previous possessors towards those individuals with whom they had shared.