Changeover cost and switching between concurrent adjusting schedules.

Using pigeon subjects we studied the effects of changeover cost on rates of switching between concurrent adjusting variable ratio schedules. Each side-key schedule while activated increased with the delivery of each reinforcer and decreased with reinforcer delivery when the alternate key was active. With no changeover cost, changeovers were frequent although not optimal. Switching from one side-key to the other was reduced following the addition of a single center-key changeover response, and reduced progressively further with increasing time (FI;FT), or response (FR;VR) requirements. This procedure is discussed as a model for the determination of variables which control a forager's giving-up in one food source patch and switching to another.

Time reallocation in a multiresponse environment: Effects of restricting response classes.

Four adult male rats were each placed for three hours daily into an apparatus that provided individual compartments for six separate location-defined responses. The available responses consisted of: (1) the opportunity to turn off room lighting, producing darkness; (2) the opportunity to view a female rat; (3) the opportunity to turn off white noise; (4) the opportunity to drink; (5) the opportunity to eat; and (6) "other," representing time in the hallway between compartments. Each subject underwent a series of conditions characterized as an A-B-A-C-A design. Manipulations consisted of the removal of a low-probability response (darkness) and of a high-probability response (escape from noise) in a counter-balanced manner across subjects. The dependent measure for all subjects was the percentage of total session time spent in each compartment. Four predictive rules concerning the redistribution of behavior after response restriction were tested, including the constant-ratio rule, equal time redistribution, the most probable alternative, and the sequential-dependency rule. The results indicate no support for any of the four predictive rules and suggest that empirical assessment of restriction effects is necessary in reinforcement studies involving temporally extended responses.

Schedule-induced locomotor activity in humans.

In two experiments, humans received tokens either on a fixed-interval schedule for plunger pulling or various response-nondependent fixed-time schedules ranging from 16 to 140 seconds. Locomotor activity such as walking, shifting weight, or pacing was recorded in quarters of the interreinforcement interval to examine the induced characteristics of that behavior in humans. While performance was variable, several characteristics were present that have counterparts in experiments with nonhumans during periodic schedules of food reinforcement: (a) first quarter rates, and sometimes overall rates, of locomotor activity were greater during intervals that terminated in a visual stimulus and token delivery than those without: (b) overall rates of locomotor activity were greater during fixed-time 16-second schedules than during fixed-time 80- or 140-second schedules; (c) rates of locomotor activity decreased during the interreinforcement intervals; (d) locomotor activity was induced by response-dependent and response-nondependent token delivery. These results showed that the rate and temporal pattern of locomotor activity can be schedule-induced in humans.

Influence of cattle age, lithium chloride dose level, and food type in the retention of food aversions.

The influences of animal age and lithium chloride (LiCl) dose levels were evaluated in forming and retaining food aversions to licorice-flavored alfalfa pellets and beet pulp. Lithium chloride (100, 200, 300 mg/kg BW) was given to yearling and mature cattle by gavage (four animals per age-dose treatment group) after meals of the test food. Cattle dosed with the higher levels of LiCl formed aversions more rapidly in both trials than those dosed at lower levels. Both age classes extinguished aversions to licorice-flavored alfalfa pellets, but aversions to beet pulp persisted. Mature cattle retained aversions to beet pulp to a greater extent than did young cattle in social facilitation and single-choice persistence trials. Of dosages studied, the optimum for retaining aversions to beet pulp was 200 mg/kg BW for mature cows. Yearling cattle required 300 mg/kg for complete abstinence.

Operant analysis of chronic locoweed intoxication in sheep.

Five sheep were fed a 10% locoweed (Oxytropis sericea) pellet or alfalfa pellets for 3- to 5-wk periods to determine the effects of intermittent locoweed ingestion on operant responding; three controls were fed alfalfa pellets for 22 wk. Sheep were trained to respond to a multiple schedule with a fixed ratio (FR) 5 and fixed interval (FI) 50 s as major elements; performance was reinforced with rolled barley. Locoweed-treated sheep decreased (P < .05) FR response rate after 4 wk of locoweed feeding, but this decrease first appeared during the first recovery period (wk 6). The FR response rate of intoxicated sheep did not return to baseline during the remainder of the study and differed from controls during most of the study. Controls did not deviate (P > .05) from their FR baseline except during wk 2. Sheep did not stabilize on the FI component. As locoweed-treated sheep became progressively more intoxicated, they altered their pattern of FR responses, with longer post-reinforcement pauses, and a slower overall FR rate. Intoxicated sheep ingested an average of .21 mg swainsonine.kg-1.d-1. Overt signs of intoxication were noted when two sheep were stressed on wk 17. These two sheep had neuroviseral vacuolation typical of locoweed poisoning, whereas the three remaining locoweed-treated sheep that were euthanatized 5 wk later showed little histologic evidence of intoxication. Our findings indicate that "on-off" or cyclic grazing of locoweed ranges should be approached cautiously. Such a grazing program may be feasible because of the rapid resolution of histologic pathology; however, an initial toxic insult of 4 wk seems to be excessive, even at low doses, because sheep may exhibit persistent behavioral abnormalities that require > 6 wk to resolve.

Conditioning taste aversions to locoweed (Oxytropis sericea) in horses.

Locoweed (Oxytropis sericea) is a serious poisoning problem for horses grazing on infested rangelands in the western United States. Our objectives were to determine 1) whether lithium chloride or apomorphine would condition aversions to palatable foods, and at what doses, and 2) whether horses could be averted to fresh locoweed in a pen and grazing situation. Apomorphine was not an acceptable aversive agent because at the dose required to condition an aversion (> or = 0.17 mg/kg BW), apomorphine induced unacceptable behavioral effects. Lithium chloride given via stomach tube at 190 mg/kg BW conditioned strong and persistent aversions to palatable feeds with minor signs of distress. Pen and grazing tests were conducted in Colorado to determine if horses could be averted to fresh locoweed. Pen tests indicated that most horses (5/6) were completely averted from locoweed. Treated horses ate 34 g of fresh locoweed compared to 135 g for controls (P < 0.01) during three pen tests when offered 150 g per test. One horse (T) in the treatment group ate locoweed each time it was offered in the pen, but ate no locoweed while grazing. In the grazing trial, control horses averaged 8.6% of bites of locoweed (P < 0.01) during the grazing portion of the study, whereas treated horses averaged <0.5%. One treated horse (S) accounted for all consumption; he consumed 15% of his bites as locoweed in a grazing bout on d 2 of the field study. Thereafter, he was dosed a second time with lithium chloride and ate no locoweed in the subsequent 5 d. Three of six horses required two pairings of lithium chloride with fresh locoweed to condition a complete aversion. The results of this study indicate that horses can be averted from locoweed using lithium chloride as an aversive agent, and this may provide a management tool to reduce the risk of intoxication for horses grazing locoweed-infested rangeland.

Effect of previous locoweed (Astragalus and Oxytropis species) intoxication on conditioned taste aversions in horses and sheep.

Locoweed species (Astragalus and Oxytropis spp.) are a serious toxic plant problem for grazing livestock. Horses and sheep have been conditioned to avoid eating locoweed using the aversive agent LiCl. The objective of this study was to determine if previous locoweed intoxication affects food aversion learning in horses and sheep. Horses and sheep were divided into 3 treatment groups: control (not fed locoweed and not averted to a novel feed); locoweed-novel feed averted (fed locoweed and averted to a novel feed); and averted (not fed locoweed and averted to a novel feed). Animals in the locoweed-novel feed averted groups were fed locoweed during 2 periods of 21 and 14 d, respectively, with each feeding period followed by a 14-d recovery period. Animals were averted to a novel test feed at the end of the first locoweed-feeding period, and periodically evaluated for the strength and persistence of the aversion. During the first recovery period, locoweed-novel feed averted horses ate less (9.5% of amount offered) of the test feed than did control horses (99.8%) and did not generally differ from averted horses (0%). During recovery period 2, locoweed-novel feed averted horses (4.3%) differed (P = 0.001) in consumption (% of offered) of the test feed from controls (100%) and the averted group (0%). Locoweed-novel feed averted sheep differed (P = 0.001) from controls (14.4 vs. 99.5%, respectively, during recovery period 1), whereas locoweed-novel feed averted sheep did not differ (P > 0.50) from averted sheep (0.6%). During the second recovery period, control sheep (100%) differed (P < 0.05) from averted (0%) and locoweed-novel feed averted (12.2%) groups. Two intoxicated sheep (locoweed-novel feed averted) partially extinguished the aversion during the first recovery period, but an additional dose of LiCl restored the aversion. Two of 3 intoxicated horses had strong aversions that persisted without extinction; 1 horse in the locoweed-novel feed averted group had a weaker aversion. These findings suggest that horses and sheep previously intoxicated by locoweeds can form strong and persistent aversions to a novel feed, but in some animals, those aversions may not be as strong as in animals that were never intoxicated.

Reversal of tall larkspur (Delphinium barbeyi) poisoning in cattle with physostigmine.

Tall larkspur poisoning of cattle is a serious problem on western US rangelands. Single oral doses of tall larkspur ranging from 1.5 to 3 g/kg body weight were administered to steers. These doses caused clinical signs of muscular tremors and collapse. Physostigmine was administered iv, ip or sc at 0.04 to 0.08 mg/kg body weight when animals were sternally or laterally recumbent. Physostigmine given iv rapidly reversed the larkspur toxicity. Serial injections of physostigmine were generally necessary to reverse acute toxicity. Administration of physostigmine to grazing animals poisoned on larkspur was also effective. Physostigmine can be effective treatment for intoxicated cattle consuming tall larkspurs.

Potential aversive compounds in leafy spurge for ruminants and rats.

Several wild and domestic ruminant species and horses apparently will not consume leafy spurge (Euphorbia esula) while grazing range and pasture lands. It has been demonstrated that leafy spurge can elicit conditioned food aversions in cattle and sheep, and the aversion-eliciting capacity of leafy spurge may account for why cattle seldom graze this nutritious plant and why sheep may not readily consume it at some locations. The identity of the aversive compound(s) in leafy spurge is unknown, but several different diterpenoid ingenol esters have been isolated from its tissues, and we suspect that one or more ingenol esters may be aversion-eliciting compounds in leafy spurge. The objectives of this study were to determine whether or not leafy spurge is aversive to laboratory rats and if a crude acetone extract of leafy spurge, presumably containing ingenol esters and other phytochemicals, could generate an aversive response in sheep and laboratory rats. An additional objective was to determine whether or not a particular ingenol monobenzoate, which may be similar to ingenol esters in leafy spurge, might also elicit an aversive response from rats. Rats exhibited food aversions associated with leafy spurge (P < 0.05). An acetone extract of leafy spurge induced conditioned food aversions in both sheep and rats (P < 0.01). The ingenol 3-monobenzoate also induced conditioned food aversions in rats (P < 0.01). Our interpretation of these data is that rats can be used as a model for cattle and sheep with respect to their aversion to leafy spurge ingestion. Additionally, we suggest that one or more ingenol esters may be aversion-inducing agents in leafy spurge. However, others may exist in leafy spurge that are also aversive or are the only or prime aversive chemicals.

Locoweed embryotoxicity in rats.

In an effort to develop a laboratory animal assay for locoweed (Astragalus lentiginosus and Astragalus wootoni) teratogenicity, different amounts of the plant were administered either in feed pellets or by gavage to rats. In preliminary experiments with A. lentiginosus in feed pellets, we found dose-dependent effects on the number of pups born alive and on birth weight of pups. In the primary experiment, 1 gm of A. lentiginosus or A. wootoni administered by gavage on Days 7 through 17 of gestation produced similar results. A. lentiginosus reduced pup birth weights by 13% less than that of controls; this weight differential persisted for at least 4 weeks. A. lentiginosus also reduced the number of offspring born alive (34.9% fewer than those of controls) and the number of survivors to weaning (86.4% fewer than that of controls). No gross malformations were observed. Behavioral tests, however, showed that activity patterns of 30-day-old offspring of dams fed locoweed differed from those of pups of control dams. A test of learning indicated no differences. At 80 days of age, offspring of treated dams showed no significant differences from those of control dams. The same behavioral tests also showed no differences among the dams at the time of testing. Microscopic examination of tissues from pups whose mothers had been given locoweed revealed lesions only in the newborn pups. Neural lesions were not observed even in the newborn pups. Dose-dependent and species-dependent embryotoxic effects of locoweed were observed in rats.

Self-organization of foraging behaviour: from simplicity to complexity without goals.

A herbivore faces challenges while foraging-ongoing changes in its physiological condition along with variation in the nutrient and toxin concentrations of foods, spatially and temporally-that make selecting a nutritious diet a vital affair. Foraging behaviours arise from simple rules that operate across levels of resolution from cells and organs to individuals and their interactions with social and physical environments. At all these levels, behaviour is a function of its consequences: a behaviour operating on the environment to induce changes is itself changed by those events. Thus, behaviour emerges from its own functioning-behaviour self-organizes-not from that of its surroundings. This ostensible autonomy notwith-standing, no self-organizing system (cell, organ, or individual) is independent of its environs because existence consists of an ongoing exchange of energy and matter. According to this view, the notion of cause and effect is replaced with functional relationships between behaviours and environmental consequences. Changes in physical environments alter the distribution, abundance, nutritional, and toxicological characteristics of plants, which affect food preference. Social interactions early in life influence behaviour in various ways: animals prefer familiar foods and environments, and they prefer to be with companions. Animals in unfamiliar environments often walk farther, ingest less food, and suffer more from malnutrition and toxicity than animals in familiar environments. An individual's food preferences-and its ability to discriminate familiar from novel foods-arise from the functional integration of sensory (smell, taste, texture) and postingestive (effects of nutrients and toxins on chemo-, osmo-, and mechano-receptors) effects. The ability to discriminate among foods is critical for survival: all problems with poisonous plants are due to an inability to discriminate or a lack of alternatives. Animals eat a variety of foods as a result of nearing or exceeding tolerance limits for sensory and postingestive effects unique to each food. After eating any food too frequently or excessively, the likelihood increases that animals will eat alternative foods owing to exceeding sensory-, nutrient-, and toxin-specific tolerance limits. Cyclic patterns of intake of a variety of foods reflect seemingly chaotic interactions among flavours, nutrients, and toxins interacting along continua.