Comparative ingestive mastication in domestic horses and cattle: a pilot investigation.

It is often assumed that horses chew food more intensively during ingestion than cattle, which - as ruminants - complete part of the mastication during rumination. This has been proposed as a reason for more robust mandibles, larger masseter insertion areas and larger masseter muscles in horses as compared to cattle and other grazing ruminants. In this study, we evaluate results of comparative feeding trials with three horses (338-629 kg) and three cows (404-786 kg), on four different roughages. Ingestion time (s/g dry matter) and chewing intensity (chews/g dry matter) differed among animals within a species, indicating an influence of body mass, and differed significantly between different forages. However, although numerical differences clearly suggest that horses have longer ingestion times and higher chewing intensities on high-fibre roughage than do cattle, this could not be proven in this dataset, most likely because of the small number of individuals sampled. Further studies are required to corroborate the suspected ingestive behaviour difference between equids and ruminants.

Crib-biting in US horses: breed predispositions and owner perceptions of aetiology.

REASONS FOR PERFORMING STUDY: Crib-biting is an equine stereotype that may result in diseases such as colic. Certain breeds and management factors have been associated. OBJECTIVES: To determine: breed prevalence of crib-biting in US horses; the likelihood that one horse learns to crib-bite from another; and owner perceptions of causal factors. METHODS: An initial postal survey queried the number and breed of crib-biting horses and if a horse began after being exposed to a horse with this habit. In a follow-up survey, a volunteer subset of owners was asked the number of affected and nonaffected horses of each breed and the extent of conspecific contact. The likelihood of crib-biting given breed and extent of contact was quantified using odds ratio (OR) and significance of the association was assessed using the Chi-squared test. RESULTS: Overall prevalence was 4.4%. Thoroughbreds were the breed most affected (133%). Approximately half of owners believed environmental factors predominantly cause the condition (54.4%) and crib-biting is learned by observation (48.8%). However, only 1.0% of horses became affected after being exposed to a crib-biter. The majority (86%) of horses was turned out in the same pasture with other horses and extent of contact with conspecifics was not statistically related to risk. CONCLUSION: This is the first study to report breed prevalence for crib-biting in US horses. Thoroughbreds were the breed more likely to be affected. More owners believed either environmental conditions were a predominant cause or a combination of genetic and environmental factors contributes to the behaviour. Only a small number of horses reportedly began to crib-bite after being exposed to an affected individual, but approximately half of owners considered it to be a learned behaviour; most owners did not isolate affected horses. POTENTIAL RELEVANCE: Genetic predisposition, not just intensive management conditions and surroundings, may be a factor in the high crib-biting prevalence in some breeds, and warrants further investigation. Little evidence exists to suggest horses learn the behaviour from other horses, and isolation may cause unnecessary stress.

Efficacy and effects of various anti-crib devices on behaviour and physiology of crib-biting horses.

REASONS FOR PERFORMING STUDY: Crib-biting is a common oral stereotypy. Although most treatments involve prevention, the efficacy of various anti-crib devices, including surgically implanted gingival rings, has thus far not been empirically tested. OBJECTIVES: Demonstrate the effect that 2 anti-crib collars, muzzle and gingival rings have on crib-biting, other maintenance behaviours, and cortisol levels in established crib-biting horses. STUDY DESIGN: Randomised, crossover clinical trial. METHODS: In Experiment I, 2 anti-crib collars and a muzzle were used on 8 established crib-biting horses; horses wore each of 3 devices for 7 days, with a 7-day device-free period between treatments. Horses were video recorded for 24 h at least 3 times each week prior to any device placement, and always the day after a device was removed. In Experiment II, gingival rings were used in 6 established crib-biting horses; horses were video recorded for 3 days prior to ring implantation and the day after surgery until the rings became ineffective. Plasma cortisol levels were assessed every day during Experiment II and on Days 1, 3 and 5 of each week during Experiment I. RESULTS: All devices significantly reduced crib-biting compared with control periods. There was no significant difference in crib-bite reduction amongst devices in Experiment I, or between pre-device periods and the first day the device was removed. The only increase in plasma cortisol occurred on the day of surgery in Experiment II. CONCLUSIONS: Common anti-crib devices are effective in reducing crib-biting and significant distress was not evident from our findings. We did not find a post inhibitory rebound effect. Surgical rings were successful only temporarily and implantation was probably painful to the horses. Because stereotypies involve suboptimal environmental conditions, efforts should be made to improve husbandry factors previously shown to contribute to crib-biting, and research into decreasing horses' motivation to crib-bite should continue.

Gastrointestinal factors in hunger and satiety.

The evidence for the participation of signals from the gastrointestinal tract in hunger and satiety is reviewed. Various methods of subjecting only certain portions of the gastrointestinal tract to food are described including sham feeding, crossing of the intestines of two animals, and intragastric, intraduodenal and intravenous feeding. These methods have revealed that animals eat more when only the mouth is exposed to food, but consume less when food enters the stomach directly and still less when feeding themselves intravenously. The role of the stomach in satiety is discussed in relation to the results of the many experiments in which intragastric loads were administered. These experiments generally revealed that gastric loads suppressed intake in proportion to their caloric value, but that compensation was not precise. A similar analysis is made of the role of the intestines in satiety. These studies present evidence for osmoreceptors and glucoreceptors, distention, and pressure, all of which may participate in satiety. The numerous effects of vagotomy on food intake are reviewed and an attempt is made to separate afferent from efferent effects and vagal effects from nonspecific surgical injury. The role of other afferents is explored. A brief survey of the most recent studies on humoral factors indicates that cholecystokinin is probably acting as a satiety agent, but its mode of action remains unknown. The changes in the gastrointestinal tract that accompany an increase or decrease in food intake reveal adaptive changes that occur in the organ. The evidence for gastrointestinal factors in satiety of the suckling animal is presented. Finally, a hypothesis is presented in which the various factors that bring a meal to an end are included. Preabsorptive satiety may consist of multiple factors which probably include signals from osmoreceptors and possibly other chemoreceptors in the upper small intestine. The signals from these receptors are neural while other products of digestion stimulate the release of humoral agents, such as cholecystokinin, which may act locally, in the liver, or on the central nervous system. Stretch or distention of the intestines at various locations may also contribute to preabsorptive satiety.

Thirst and salt appetite in horses treated with furosemide.

When a preliminary experiment in sodium-replete ponies revealed an increase, but not a significant increase, in salt consumption after furosemide treatment, the experiment was repeated using sodium-deficient horses in which aldosterone levels might be expected to be elevated to test the hypothesis that a background of aldosterone is necessary for salt appetite. Ten Standardbred mares were injected intravenously with furosemide or an equivalent volume of 0.9% sodium chloride as a control to test the effect of furosemide on their salt appetite and blood constituents. Sodium intake and sodium loss in urine, as well as water intake and urine output, were measured and compared to determine accuracy of compensation for natriuresis and diuresis. Plasma protein and packed cell volume showed significant increases in response to furosemide treatment (F = 29.31, P less than 0.001 and F = 11.20, P less than 0.001, respectively). There were no significant changes in plasma sodium concentration or osmolality in response to the treatment (P greater than 0.05). The furosemide-treated horses consumed 126 +/- 14.8 g salt, significantly more than when they were given the control injection (94.5 +/- 9.8 g; t = 2.22, P = 0.05). In response to furosemide, horses lost 962 +/- 79.7 and consumed 2,170 +/- 5 meq sodium; however, compared with control, they lost 955 meq more sodium and ingested only 570 meq more sodium, so they were undercompensating for natriuresis. The furosemide-treated horses drank 9.6 +/- 0.8 kg of water, significantly more than when they received the control injection (6.4 +/- 0.8 kg; t = 6.9, P less than 0.001).(ABSTRACT TRUNCATED AT 250 WORDS)

Comparison of cerebrospinal fluid monoamine metabolite levels in dominant-aggressive and non-aggressive dogs.

Aggression has been shown to be related to reduced serotonergic activity in humans and non-human primates, and in rodents. We now studied the relationship between cerebrospinal fluid (CSF) monoamine metabolites and canine aggression in 21 dominant-aggressive dogs (Canis familiaris) and 19 controls. The diagnosis of dominance-related aggression was based upon a history of biting family members in contexts associated with dominance challenges. Post-mortem CSF 5-HIAA, MHPG and HVA were measured by high-performance liquid chromatography using electrochemical detection. Concentrations of CSF 5-HIAA (P = 0.01) and HVA (P < 0.001) were lower in the aggressive group (median values: 5-HIAA 202.0 pmol/ml; HVA 318.0 pmol/ml) than in controls (5-HIAA 298.0 pmol/ml; HVA 552.0 pmol/ml). No differences were noted in CSF MHPG levels. Differences in 5-HIAA were maintained after controlling for breed and age of dogs, but HVA differences may have been breed-dependent. Lower levels of 5-HIAA (P = 0.02) and HVA (P = 0.04) were found in the subgroup of aggressive dogs with a history of biting without warning (5-HIAA 196.0 pmol/ml; HVA 302.0 pmol/ml) compared to dogs that warned (5-HIAA 244.0 pmol/ml; HVA 400.0 pmol/ml). This study suggests that reduced serotonergic function is associated with aggressive behavior and impaired impulse control in dogs, a finding that is consistent with observations in primates, and suggests that serotonin modulates aggressive behavior throughout mammals.

The role of the Flehmen response in the behavioral repertoire of the stallion.

The role of the Flehmen response in equine behavior was investigated under field and laboratory conditions. In Experiment 1, a field study made of five stallions on pasture with between three and eighteen mares each during the season indicated the following: 1) The Flehmen response was most frequently preceded by nasal, rather than oral, investigation of substances; 2) The stallions' rate of Flehmen varied with the estrous cycles of the mares; 3) The rate of Flehmen response did not show a variation with time of day; and 4) The Flehmen response was most frequently followed by marking behaviors rather than courtship behaviors. The results suggest that the Flehmen response is not an immediate component of sexual behavior, e.g., courtship of the stallion but may be involved in the overall monitoring of the mare's estrous cycle. Therefore the Flehmen response may contribute to the chemosensory priming of the stallion for reproduction. In Experiment 2 stallions were presented with urine or feces of mares in various stages of the reproductive cycle as well as with their own or other males' urine or feces. The occurrence of sniffing and Flehmen was used to determine the discriminatory ability of the stallions. Stallions can differentiate the sex of a horse on the basis of its feces alone, but cannot differentiate on the basis of urine. This ability may explain the function of fecal marking behavior of stallions.

Gastric emptying and cholecystokinin in the control of food intake in suckling rats.

The rate of gastric emptying was determined in three-day old suckling rats. The gastric loads were given by gavage, and, after from 5 to 100 min, emptying was determined by removing the stomach and weighing the contents. Results were expressed as percentage of load still remaining in the stomach at one hour. The gastric loads in increasing order of speed of emptying were 1.0 M Na acetate, heavy cream, 0.5 M NaCl, milk, corn oil, 0.15 M lactose, 0.3 M glucose, 0.15 M NaCl, acidic water, and water. The rate of emptying was compared to the effectiveness in previous experiments of the same gastric loads in depressing intake. There was no significant correlation between rate of gastric emptying of the loads and their effectiveness in producing satiety. The octapeptide of cholecystokinin (80 Ivy dog units or 2.7 micrograms/kg i.p.) significantly depressed intake (measured as weight gain) of suckling rats of 1 1/2 hours, but the same dose did not slow gastric emptying. These findings indicate that rate of gastric emptying does not determine satiety in the suckling rat.

Vasopressin in dehydrated and rehydrated ponies.

Six pony mares deprived of water for 24 hours showed significant increases in plasma vasopressin (2.8 pg/ml) and osmolality (9 mosmol/kg). When water was made available the ponies drank rapidly (5 of 6 drank to satiety within 90 seconds) and corrected their fluid deficits precisely. Vasopressin did not return to predehydration levels until osmolality did after 15 minutes of access to water. The horse differs from rodents and humans, but is similar to pigs in that vasopressin levels do not fall before osmolality returns to normal. Oropharyngeal factors, therefore, may not be as important in vasopressin release in horses as in other species.

Changes in free fatty acids and triiodothyronine in response to feeding in pigs.

Ten pigs were surgically implanted with jugular venous catheters. Blood samples were acquired before, during and after meals that occurred ad lib or after 5-hr or 17-hr fasts. 3,3',5-Triiodothyronine (T3) concentration increased (p less than 0.05) from 0.49 +/- 0.04 to 0.91 +/- 0.13 ng/ml (mean +/- SE) following feeding only when the pigs had been fasted for 17 hr prior to the meal. In contrast, free fatty acid levels decreased (p less than 0.05) with feeding under ad lib, 5-hr fasted and 17-hr fasted conditions. Free fatty acid concentration decreased from 222 +/- 66 mEq/l to 91 +/- 20 mEq/l (mean +/- SE) in pigs fed ad lib. Free fatty acids may function as hunger signals or their decrease may serve as a satiety signal in free-feeding animals. T3 may buffer against a large caloric intake associated with large meals, by increasing dietary thermogenesis.

Stimuli of thirst in donkeys (Equus asinus).

A study of the stimuli of thirst was conducted on six feral donkeys. Donkeys were found to be stimulated to drink by overnight water deprivation, by the diuretic furosemide, and by hypertonic saline infusion, all in the absence of heat stress or work. Donkeys compensate accurately for the fluid deficit caused by overnight water deprivation. After 19 hr without water, they drank 8.8 +/- 2.4 (mean +/- SE) liters within 60 min. Their undeprived overnight intake was 8.4 +/- 1.5 liters. However, latency was longer and water intake was less than that of ponies with the same changes in blood parameters, suggesting that donkeys have a higher thirst threshold than ponies. Further, plasma volume fell less in donkeys, but osmotic changes were similar to those reported in ponies exposed to the same deprivation. Donkeys infused with 250 ml of 15% NaCl drank 0.7 +/- 0.6 liters of water within 45 min, and osmolality increased from 287 to 297 mosmol/kg water; they drank no water in the same time period when infused with 250 ml 0.9% NaCl (p less than 0.05). Donkeys injected IV with 2 mg/kg furosemide drank 3.8 +/- 1.1 liters within 3 hr. Plasma protein increased from 6.9 to 7.8 g/dl. When injected with 0.9% NaCl they drank 1.0 +/- 0.5 liters (p less than 0.05). In sum, the positive thirst responses of these donkeys to cellular and extracellular dehydration were similar to those earlier demonstrated in ponies, but the results suggest a less sensitive response, albeit combined with a better internal defense of blood volume.

Effects of duodenal osmotic loads on spontaneous meals in pigs.

Fourteen young Large White pigs were trained to press a panel switch to obtain reinforcements of food (FR 10-15). Duodenal injections (5 ml/kg) were made automatically via implanted catheters shortly after the beginning of every other meal 24 hours a day. Solutions of varying osmoconcentration and nutrient content were injected. Isosmotic solutions of glucose (5, 20 and 40%) and NaCl (0.9, 3.25 and 6.5%) equally depressed the size of ongoing meals in proportion to their hypertonicity. For example, 20% glucose and 3.25% NaCl depressed meal size to 62% of control, and 40% glucose and 6.5% NaCl to 34%. Xylose (33%) was less effective than its glucose osmotic equivalent. Tetracaine (0.5%) blocked most of the reduction of meal size. The reduced meal size was due to a decrease in meal duration. Neither rate of eating during the meal nor intermeal interval changed. The results support the hypothesis that a duodenal osmoreceptive system participates normally in the control of meal size.

The effects of caloric dilution on meal patterns and food intake of ponies.

In order to determine if horses will increase their intake in response to caloric dilution, four pony geldings were fed ad lib a mixed grain diet either undiluted (3.4 Mcal/kg of digestible energy) or diluted (wt/wt) with 25% sawdust (2.6 Mcal/kg) or with 50% sawdust (1.7 Mcal/kg). The mean daily caloric intake was 17,457 kcal (3.4 Mcal diet), 17,546 kcal (2.6 Mcal diet) and 12,844 kcal (1.7 Mcal). The mean time spent eating was 246 (3.4 Mcal), 351 (2.6 Mcal), and 408 (1.7 Mcal) minutes/day. Meal size increased and meal frequency decreased with increasing dilution. The median long survivorships of intermeal intervals were 6.4 min (3.4 Mcal), 3.95 min (2.6 Mcal) and 4.91 min (1.7 Mcal). Ponies responded to caloric dilution by increasing the volume of intake to maintain caloric intake when the diet had 25% diluent. When the diet was diluted by 50%, intake was increased, but not at a rate adequate to maintain caloric intake. However, the ponies were able to maintain body weight.

Friendliness to humans and defensive aggression in cats: the influence of handling and paternity.

This study was undertaken to examine and reduce the stress and aggressiveness associated with fear of handling in laboratory cats (Felis sylvestris catus). Thirteen litters of kittens from a specific pathogen-free breeding colony were divided into three treatment groups: two were early weaned, removed from the colony and caged individually with or without handling up to 8 weeks of age, and the third served as a control group, removed from the colony just before testing. Behavior tests measuring degree of friendliness to humans and response to physical restraint were performed at ages 8, 12, 16, and 20 weeks. Serum cortisol concentrations were measured after each test. Results indicate that litter and sire influenced tractability but that handling or individual caging of kittens did not. Posttest serum cortisol concentrations were below normal adult levels in most kittens, including those reacting fearfully during testing and aggressively during restraint, and, therefore, are not a reliable indicator of stress in juvenile cats.

Effect of meal frequency on fluid balance and behavior of ponies.

Twelve ponies were fed their total daily ration either as one large meal or divided into six small meals. Pre- and post-feeding behavior was recorded six times a day. Blood samples were taken for 30 min before and two hr after the meal. Plasma protein increased from 7.0 to a peak of 7.3 g/dl with small meals and from 7.3 to 8.1 g/dl with large meals, and returned to pre-feeding levels by 90 min post-feeding. Hematocrit rose from 33.3 to 34.1% with small meals and from 33.0 to 36.0% with large meals. These rapid and short-lived increases indicate a decrease in plasma volume. Plasma osmolality rose with feeding from 283 to 285 mosmoles/kg with small meals and from 281 to 288 mosmoles/kg with large meals. Water availability had no significant effect on blood changes. Digestibility and rate of passage were measured with chromic oxide, but there were no differences. Vocalizing (neighing) and walking occurred more often before than after feeding, while eating bedding and engaging in other oral behaviors were more frequent after feeding.

Glucoprivic (2DG) eating in rats despite knife cut induced hyperphagia.

Two weeks after parasagittal hypothalamic knife cuts, baseline eating was elevated and 300 mg/kg 2 deoxy-D-glucose (2DG) did not further stimulate food intake. Five weeks postoperatively the food intake baseline had fallen and an eating response to 300 mg/kg 2DG was now seen (p less than 0.005). In this delayed (static) phase intake was also stimulated by 150 mg/kg 2DG (p less than 0.005). 600 mg/kg did not stimulate intake in the lesioned rats at any time, although sham-operated rats always responded positively to this high dose. In conclusion, the neural substrate damaged in hypothalamic hyperphagic rats does not appear to mediate eating in response to glucopribation. The eating response is masked by high baseline intake in the dynamic phase, but reappears in the static phase.

Stimulation of food intake in horses by diazepam and promazine.

In two adult horses doses of 0.02-0.03 mg/kg diazepam, intravenously, increased 1 hr intake 54-75% above control levels. Intake was stimulated when the diet was a high grain, calorically dense one and also when the diet was a high fiber, calorically dilute one. Two young rapidly growing weanling horses showed an even more pronounced stimulation of intake. Following diazepam 1 hr intake was increased 105-240% above control lelvels. Promazine at a dose of 0.5 mg/kg also stimulated intake in adult horses, but not as markedly as did diazepam. A transquilizer and a neuroleptic appear to have a stimulatory eff upon short-term intake in horses.

The flehmen response of bulls and cows.

The response of dairy bulls to the urine of cows in various stages of the reproductive cycle was quantified by presenting 200 ml of urine in a stainless steel bowl to the stanchioned bulls for 10 min. Estrous mucus was also presented in the same manner. Sniffs, nose licks and flehmen responses were recorded. Of 15 bulls tested only 7, or 47%, met the criteria of two flehmens in response to estrous urine. Among these bulls, the rate of flehmen was higher to estrous urine 6.1 +/- 1 flehmen/10 min) than to nonestrous urine (3.5 +/- 0.6 flehmen/10 in, paired t = 3.1, P < 0.03). Flehmen duration was also longer in response to estrous urine (6.4 +/- 0.4 sec) than to nonestrous urine (5.7 +/- 0.4 sec, t = 2.65, P < 0.03). There were no significant differences between the sniff frequencies and durations or in the number of licks to estrous and diestrous urine. There were significantly more flehmen responses to estrous urine (7 +/- 1.4 10 min ) than to mucus (2.25 +/- 0.9 10 min , t = 4.75, P < 0.01). The response to water (0.6 +/- 0.3 10 min ) was not different from that to mucus (t = 2.37, P < 0.10). The spontaneous flehmen rate of dairy bulls in their home stalls was 3.2 +/- 0.7 24 h . Although estrous cows did not exhibit flehmen frequently, they did sniff bull urine more frequently (3.6 +/- 0.6 10 min ) than nonestrous cows (1.8 +/- 0.3 10 min , t = 2.4, P < 0.03).

Stability of equine hierarchies and the prevention of dominance related aggression.

The dominance hierarchy of a herd of 10 Thoroughbred mares was determined twice, at an interval of 18 months, using paired feeding tests. Each mare's rank was correlated significantly between the 2 tests. This indicated that the hierarchy within the herd was stable. The offspring of dominant and subordinate mares were also tested for dominance in their own age groups. The offspring of dominant mares tended to be near the top of the hierarchy while those of middle and low ranking mares were not consistently found in the middle or bottom of their own hierarchies. Paired feeding tests were carried out on 8 ponies. During tests the time that each pony spent eating and the ponies' aggressive interactions were recorded. Two situations were used. Each pony-pair was tested when both ponies were in the same paddock and also when they were separated by a rail fence. The subordinate ponies spent significantly more time eating and the domonant pony was significantly less aggressive, when the pony-pair was separated by a fence than when they were in one paddock. It was concluded that the dominance hierarchies of adult horse groups changed very little over time and that the foals of dominant mares will tend to be dominant in their own age groups. Management practices can be used to reduce aggression and consequent injury that may arise in group feeding situations.

Coprophagy by foals: effect of age and possible functions.

In colts and fillies observed from birth to 24 weeks old, coprophagy occurred from Weeks 1 to 19. Its frequency was greatest during the first two months. Coprophagy was rarely observed in mares and stallions. Foals usually ate the faeces of their mother but were observed to eat their own and those of a stallion and another unrelated mare. Urination by the foal occurred before, during or after 26 per cent of the coprophagy incidents. It is hypothesised that foals may consume faeces in response to a maternal pheromone which signals the presence of deoxycholic acid or other acids which the foal may be deficient in and which it may require for gut immuno-competence myelination of the nervous system. Such a pheromone may also serve to accelerate growth and sexual maturation. Coprophagy may also provide nutrients and introduce normal bacterial flora to the gut.