Categorical counting

Pigeons pecked on three keys, responses to one of which could be reinforced after a few pecks, to a second key after a somewhat larger number of pecks, and to a third key after the maximum pecking requirement. The values of the pecking requirements and the proportion of trials ending with reinforcement were varied. Transits among the keys were an orderly function of peck number, and showed approximately proportional changes with changes in the pecking requirements, consistent with Weber's law. Standard deviations of the switch points between successive keys increased more slowly within a condition than across conditions. Changes in reinforcement probability produced changes in the location of the psychometric functions that were consistent with models of timing. Analyses of the number of pecks emitted and the duration of the pecking sequences demonstrated that peck number was the primary determinant of choice, but that passage of time also played some role. We capture the basic results with a standard model of counting, which we qualify to account for the secondary experiments.     

Relationship between feather pecking and ground pecking in laying hens and the effect of group size

The aim of this experiment was to study the relationship between feather pecking and ground pecking in laying hens and the effect of group size on feather pecking behaviour. Hisex White hens were kept in floor pens in group sizes of 15, 30, 60 and 120 birds, each with four replicates. Behavioural observations were performed at four different ages and focused on the number of feather pecks and aggressive pecks, both given and received. The part of the body pecked and the location of the bird was recorded as well as the number of pecks made to the floor, feeder and drinker.The results showed that most feather pecking activity occurred in the largest group size (120 birds) and there was some evidence of an increasing frequency of aggressive pecks with increasing group size. The parts of the body which were targets for feather pecking varied depending on the location of the bird giving the peck and the bird receiving it. When looking at the behaviour of individuals, birds doing a lot of feather pecking also showed more ground pecking.     

Variation of pecking rate with sward height in wild wigeon Anas penelope

We examined how pecking rate changed with sward height in wintering European wigeon Anas penelope in a nature reserve of the French Atlantic coast. Pecking rates were recorded as the time taken to perform 10 or 50 pecks. We found a negative correlation between pecking rate and sward height over a range of heights from 1.0 to 3.5 cm, but no sex effect. Equations based on the time taken to perform 10 or 50 pecks were slightly different, suggesting that scanning (scans of the surroundings) and moving (search for plant items of good quality) had an effect on pecking rate. However, these costs were lower (i.e. a decrease of 6–10% of pecking rate) than the effect of handling time, which remained the dominant foraging process constraining pecking rate (i.e. a decrease of 24% with each 1-cm increase in sward height).     

An Experimental Analysis of Spatial Position Effects on Foraging and Vigilance in Brown-Headed Cowbird flocks

Several observational studies have found that the costs and benefits of social foraging vary as a function of spatial position in the group. However, it is difficult to make mechanistic inferences because several confounding factors, such as food deprivation levels, food availability, neighbor distance, and group size can mask or amplify spatial position effects. We attempted to address experimentally the effect of spatial position on foraging and vigilance in a group, controlling for many confounding factors. We used enclosures that restricted physical but not visual interactions between brown-headed cowbirds and manipulated spatial position, flock size, and neighbor distance. Pecking rate (number of pecks per trial duration) was not related with position, but instantaneous pecking rate (number of pecks per foraging bout duration) was higher at the edge. The proportion of time spent head-up (scanning and food-handling) was also higher at the edge. For pecking rate and proportion of time spent scanning, changes in neighbor distance influenced the behavior of edge birds to a lesser extent than central birds. These results suggest that cowbirds at the edge perceived greater predation risk and that during the limited foraging time available, edge birds tried to compensate by foraging at a faster rate.     

Effect of manipulating feathers of laying hens on the incidence of feather pecking and cannibalism

Feather pecking is a problem in commercial laying hens, particularly in loose-housing systems, where many hens can be affected by only a few feather peckers. In addition, feather pecking can become an even larger problem if it spreads throughout the flock. There are several possible ways that feather pecking may spread. The simplest way is that one hen may damage the feathers of a hen, and another hen may find the damaged feathers an attractive pecking target. The aim of this experiment was to determine if damaged feathers were feather-pecked more than undamaged feathers on the same body area, and to determine whether some types of feather-body area manipulations were preferred over others as pecking stimuli. Manipulations involved damaging the feathers on the rump, tail or belly of different hens, with two or three levels of severity of manipulation at each body area. Sixteen groups of 11 Lohmann Brown hens between 26 and 28 weeks were observed with the recipient, the feather pecker and the body area that was pecked all being recorded. The feather pecks were classified separately as either gentle or severe. Damaged feathers received significantly more severe feather pecks than undamaged feathers. There were also more gentle feather pecks to damaged feathers, although this did not reach statistical significance. The feather-body area manipulations that received the greatest number of severe feather pecks were the tail feathers when they were cut very short, the rump feathers when they were trimmed, and the rump when feathers were removed. These results support the suggestion that feather pecking does indeed spread through flocks by damaged feathers becoming an attractive target for feather-pecking behaviour. An unexpected result of performing the feather manipulations was an outbreak of cannibalism in half of the experimental groups. Even though there was no visible damage to the skin of the hens after having the feathers manipulated, 13 of the 16 attacked hens were wounded on the part of the body where the feathers had been damaged in some way.     

Pecking: another measurement for filial attachment in group-reared domestic fowl chicks

This study examines different types of attachment measurements and pecking behavior in group-reared domestic chicks imprinted to a silent imprinting object. Immediately upon arriving from the hatchery, groups of heavy strain chicks were imprinted to a colored foam rubber ball for 72 h. The bond between these chicks and the imprinting object was then tested from 8 days until 5 weeks of age. Three days of imprinting sessions demonstrated that a strong attachment could be formed between the imprinted chicks and the imprinting object. In both experimental and control groups, the number of pecks was found to be significantly correlated with the latency measurements. It is, therefore, reasonable to consider the number of pecks of the imprinting object as another measurement for filial attachment.     

Comparisons of damaging feather pecking and time budgets in male and female turkeys of a traditional breed and a genetically selected male line

Feather and skin pecking leading to feather loss and tissue damage is a welfare problem in commercial turkeys. A factorial experiment was designed to compare a line of unselected turkeys that previously did not exhibit this behaviour with a genetically selected male line with a propensity for damaging pecking. We also compared the time budgets of male and female turkeys from the two lines at 3, 6 and 9 weeks of age and at different times of the day. Damaging feather pecking occurred in 32% of male and 15% of female male line turkeys but was not observed in the traditional line. Time budgets of males and females were similar. Preening increased and resting, feeding and general pecking declined with age. Resting was higher in the afternoon than in the morning and male line turkeys were observed resting more often than traditional turkeys at 3 and 6 weeks. Traditional turkeys displayed more gentle feather pecks than male line poults at 9 weeks. Male turkeys of both lines showed more strong feather pecks and pulls at 3 weeks whereas female turkeys showed more at 9 weeks of age. The frequency of strong feather pulls recorded electronically increased with age but was not related to mortality. It is suggested that damaging feather pecking in turkeys may be the result of vigorous investigative pecking.     

Cardioprotective and antioxidant effects of apomorphine

Apomorphine is a potent antioxidant that infiltrates through biological membranes. We studied the effect of apomorphine (2 microM) on myocardial ischemic-reperfusion injury in the isolated rat heart. Since iron and copper ions (mediators in formation of oxygen-derived free radicals) are released during myocardial reperfusion, apomorphine interaction with iron and copper and its ability to prevent copper-induced ascorbate oxidation were studied. Apomorphine perfused before ischemia or at the commencement of reperfusion demonstrated enhanced restoration of hemodynamic function (i.e. recovery of the work index (LVDP x HR) was 69.2 +/- 4.0% with apomorphine pre-ischemic regimen vs. 43.4 +/- 9.01% in control hearts, p < 0.01, and 76.3 +/- 8.0% with apomorphine reperfusion regimen vs. 30.4 +/- 11.1% in controls, p < 0.001). This was accompanied by decreased release of proteins in the effluent and improved coronary flow recovery in hearts treated with apomorphine after the ischemia. Apomorphine forms stable complexes with copper and with iron, and inhibits the copper-induced ascorbate oxidation. It is suggested that these iron and copper chelating properties and the redox-inactive chelates formed by transition metals and apomorphine play an essential role in post-ischemic cardioprotection.     

Predisposed visual memory of shapes in quail chicks

Newly hatched chicks will spontaneously peck at conspicuous objects in their field of view, and soon learn to distinguish between edible food particles and unpleasant tasting objects. To examine whether the selective pecking is based on the ability to memorize shapes, we analyzed pecking behavior of 1- to 2-days-old quail chicks (Coturnix japonica) by using ball- and triangle-shaped beads both painted in green. Repeated presentation of dry bead (either ball or triangle) resulted in a progressively fewer number of pecks (habituation). When chicks were tested by triangle after repeated presentation of ball, chicks showed a significant increase in the number of pecks at the triangle (dishabituation). On the other hand, when tested by ball after a series of triangle presentations, pecking frequency did not increase (no dishabituation). Chicks thus distinguished the triangle as a novel object after being habituated to ball, but did not respond to the ball after triangle. A similar asymmetry was found in one-trial passive avoidance task. Chicks were pre-trained by water-coated (neutral) triangle and then trained by methylanthranilate-coated (aversive) ball. In this case, most chicks learned to avoid the ball, and half of these successful learners pecked at the triangle; they distinguished triangle from ball. When chicks were pre-trained by neutral ball and trained by aversive triangle, on the other hand, most chicks did not distinguish the ball from triangle, and showed a generalized avoidance for both beads. Chicks may be innately predisposed to memorize a limited category of shapes such as ball, and associate them with selective avoidance.     

Effects of chronic antidepressant treatment on nigrostriatal and mesolimbic dopamine autoreceptors in the rat

Dopamine autoreceptors were studied by determining the effects of chronic antidepressant treatment on the ability of several doses of apomorphine to decrease 3,4-dihydroxyphenylalanine accumulation (an index of dopamine synthesis in vivo) after saline or γ-hydroxybutyric acid lactone (γ-butyrolactone). 3,4-Dihydroxyphenylalanine accumulation was measured in nigrostriatal [nucleus caudatus putamen] and mesolimbic [nucleus accumbens and tuberculum olfactorium] nerve terminals. Apomorphine decreased 3,4-dihydroxyphenylalanine accumulation in the nucleus caudatus putamen, tuberculum olfactorium and nucleus accumbens in a dose-related manner. Chronic imipramine (10 days) treatment attenuated the low and high dose apomorphine-induced decrease in 3,4-dihydroxyphenylalanine accumulation in the nucleus caudatus putamen to a greater extent than the tuberculum olfactorium or nucleus accumbens. In γ-butyrolactone-treated animals chronic treatment with imipramine, amitriptyline or bupropion (10 days) attenuated the low dose apomorphine effect in the nucleus caudatus putamen, but not the tuberculum olfactorium or nucleus accumbens. Only 2 days of imipramine treatment had no effect on the apomorphine-induced decrease in 3,4-dihydroxyphenylalanine accumulation in the nucleus caudatus putamen with or without γ-butyrolactone treatment. These data suggest that chronic treatment with three antidepressants produces dopamine autoreceptor subsensitivity in nigrostriatal neurons more than mesolimbic neurons and that this effect is not seen with short-term imipramine treatment.     

Neurochemical evidence for denervation supersensitivity: the effect of unilateral substantia nigra lesions on apomorphine-induced increases in neostriatal acetylcholine levels

Unilateral lesions of the nigro-neostriatal dopaminergic projection were induced by injections of 6-hydroxydopamine into the zona compacta of the substantia nigra. This resulted in a reduction of neostriatal dopamine to less than 6 percent of the control side. Two months later intraperitoneal injections of apomorphine (1 mg/kg) produced contralateral turning and a significant increase in neostriatal acetylcholine levels. The increase was significantly greater in the neostriatum ipsilateral to the lesion than in the intact side. Haloperidol (1 mg/kg) produced a significant decrease in neostriatal acetylcholine but this decrease did not differ between the “denervated” and intact neostriata. The nigral 6-hydroxydopamine lesions did not by themselves affect neostriatal acetylcholine levels. The fact that apomorphine produces a greater increase in neostriatal acetylcholine after lesions of the dopaminergic nigro-neostriatal projection supports earlier behavioral data suggestive of denervation supersensitivity of neostriatal dopaminergic receptors after these lesions.     

Cardioprotective and Antioxidant Effects of Apomorphine

Apomorphine is a potent antioxidant that infiltrates through biological membranes. We studied the effect of apomorphine (2?μM) on myocardial ischemic-reperfusion injury in the isolated rat heart. Since iron and copper ions (mediators in formation of oxygen-derived free radicals) are released during myocardial reperfusion, apomorphine interaction with iron and copper and its ability to prevent copper-induced ascorbate oxidation were studied. Apomorphine perfused before ischemia or at the commencement of reperfusion demonstrated enhanced restoration of hemodynamic function (i.e. recovery of the work index (LVDP?×?HR) was 69.2±4.0% with apomorphine pre-ischemic regimen vs. 43.4±9.01% in control hearts, p<0.01, and 76.3±8.0% with apomorphine reperfusion regimen vs. 30.4±11.1% in controls, p<0.001). This was accompanied by decreased release of proteins in the effluent and improved coronary flow recovery in hearts treated with apomorphine after the ischemia. Apomorphine forms stable complexes with copper and with iron, and inhibits the copper-induced ascorbate oxidation. It is suggested that these iron and copper chelating properties and the redox-inactive chelates formed by transition metals and apomorphine play an essential role in post-ischemic cardioprotection.     

Conditioning the pecking motions of pigeons

The spatio-temporal courses of head and neck motions of pigeons while pecking at small grains are described. Single and serial pecks are distinguished but the inter- and intraindividual variability of the peck kinetics is stressed. Pigeons were then trained with instrumental conditioning procedures to speed-up their pecking. A partial reinforcement schedule where pigeons had to peck repeatedly before receiving reward led to a mild shortening of inter-peck intervals at lower reinforcement rates but surprisingly, a lengthening at higher rates. A schedule where short inter-peck intervals were differentially rewarded yielded a pronounced abbreviation of the inter-peck intervals, but this was achieved by a reduction of the movement path rather than an increase in motion velocity. A schedule whereby increased approach velocities were differentially rewarded yielded marked movement accelerations. When pigeons were rewarded for diminished approach speeds they also showed significant movement decelerations. Finally, it is shown that pigeons could learn to reliably abort their peck approach movement when a visual stimulus signalling a penalty was occasionally presented during the approach movement. The proportion of successful peck interruptions decreased as these interruption signals occurred later during the approach phase. It is concluded that the pecking of pigeons is neither an innately fixed nor a visually ballistic movement. It is instead a multiply controlled and flexibly adaptable response pattern.     

Fast cortical oscillation after thalamic degeneration: pivotal role of NMDA receptor

We examined electrophysiological and molecular changes of the thalamocortical system after thalamic degeneration in Purkinje cell degeneration (pcd) mice. In pcd mice, neurons in specific thalamic nuclei including the ventral medial geniculate nucleus began to degenerate around postnatal day 50, whereas the visual thalamic nucleus and nonspecific thalamic nuclei remained almost intact. In association with the morphological changes, auditory evoked potentials in the primary auditory cortex (AC) began to decrease gradually. Fast Fourier transform analysis of spontaneous cortical field potentials revealed that fast oscillation (FO) around 25 Hz occurred in the AC but not in the visual cortex. Quantitative mRNA analysis demonstrated that expression of the N-methyl-D-aspartate (NMDA) receptor was up-regulated in the AC but not in the visual cortex. Systemic administration of an NMDA antagonist abolished the FO in the AC. These results indicate that increased NMDA activity may cause the FO in the AC of pcd mice.     

Visual memory of shapes in quail chicks: discrimination among 2-dimensional objects

Newly hatched chicks spontaneously peck at conspicuous objects, and soon learn to discriminate between edible food particles and inedible objects. To examine whether this discrimination is based on a chick's ability to memorize objects by shape cues, we analyzed the pecking behavior. One- to 3-day old quail chicks (Coturnix japonica) were presented with dry objects of different shapes (ball, disk, triangle and T-shape) of similar size (4 mm) and color (green). Habituation occurred after repeated presentation of any one of these objects (duration: 30 sec; interval: 4 min). When chicks showed significantly more pecks at a novel object (dishabituation), we assumed that chicks had memorized the habituated shapes and distinguished the novel object. Chicks did not show dishabituation between a ball and a disk. On the other hand, chicks discriminated a triangle or T-shape from the memorized image of disk, but did not memorize either triangle or T-shape by its shape. Similarly, chicks did not memorize the size of disks as a reference for subsequent pecking behavior. Chicks proved to have a limited ability to memorize shape and size cues for selective pecking behavior, in strong contrast to their accurate memorization of colors.     

Measurement of spontaneous motor activity by using the vibrator response method in rats

To investigate the usefulness of spontaneous motor activity (SMA) measurement using the vibrator response method, the acute effects of drugs on SMA were observed in Sprague-Dawley male rats. There were no significant differences between four devices. Methamphetamine (0.3-1 mg/kg, i.p.) and 1-2 mg/kg (s.c.) of apomorphine increased the SMA, but 0.05-0.2 mg/kg (s.c.) of apomorphine decreased the SMA. Apomorphine at 1-2 mg/kg (s.c.) significantly increased the SMA when the vibrator response method was used as compared with the Animex method. These results suggest that the vibrator response apparatus is useful for the measurement of SMA in rats.     

A probabilistic model of the spatial patterning of pecking in birds : Pilot study with young chicks

The pecking behaviour of young chicks ($\text{Gallus gallus}$) is studied in a situation involving several equivalent targets (mealworms). The question is raised whether the successive pecks are randomly distributed or whether they follow a systematic order based on the spatial arrangement of the targets. Data collected with one-week-old chicks indicate that pecking is compatible with a probabilistic model where the probability to peck at a given place is inversely proportional to the energy used for this particular peck. Pecking by chicks is interpreted as a functional compromise between random sampling and optimal exploitation of the environment.     

Subchronic methamphetamine treatment selectively attenuates apomorphine-induced decrease in 3,4-dihydroxyphenylacetic acid level in mesolimbic dopaminergic regions

To investigate mechanisms of behavioral enhancement produced by repeated doses of amphetamines, the effects of apomorphine on 3,4-dihydroxyphenylacetic acid (DOPAC) and dopamine (DA) levels were examined in various brain regions of the rat on the 4th day of withdrawal after repeated administration of saline or methamphetamine (3 mg/kg, s.c.) twice daily for 14 days. Apomorphine (0.1 and 1.0 mg/kg, i.p.) produced a dose-dependent decrease in DOPAC levels and no effect on DA levels in the olfactory tubercle, nucleus accumbens, striatum, frontal and cingulate cortices of saline-treated animals. A decrease in DOPAC levels produced by a low dose of apomorphine was attenuated selectively in the olfactory tubercle and nucleus accumbens of methamphetamine-treated animals. A high dose of apomorphine produced a significant decrease in DOPAC levels in both regions. No such attenuation was obtained in the striatum and the frontal and cingulate cortices.These results suggest that subchronic methamphetamine may induce development of hyposensitivity of presynaptic DA receptors in the mesolimbic regions, which contribute to the behavioral enhancement produced by the drug.     

Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesticus)

A selection experiment was initiated in 1996 in which selection for (HP line) and against (LP line) feather pecking was performed. The foundation stock was a White Leghorn layer strain established in 1970 and maintained since then as a random bred control line at the Institute. Six hatches were produced over three generations. At the age of 68 weeks (generation 0, 1996), 35 weeks (generation 1, 1997), 30 weeks (generation 2, 1998), and 27 weeks (generation 3, 1999) female birds were transferred to observation pens and their feather pecking behaviour was recorded. In each generation, 30 females and 8 males were selected from approximately 200 females and 60 males. The selection criterion was breeding value estimated by animal model on the trait 'number of bouts of feather pecking per bird per hour'.Feather pecking behaviour in adult hens was significantly higher in HP than in LP. In generation 2 the following was recorded: 3.10 versus 1.37 bouts per bird per hour (P<0.01), 7.04 versus 3.58 pecks per bird per hour (P<0.05) and the proportion of hens recorded feather pecking in the 180min observation period was 67 versus 56% (P<0.05). In generation 3 the following was recorded: 4.56 versus 0.63 bouts per bird per hour (P<0.001), 13.9 versus 2.51 pecks per bird per hour (P<0.001) and the proportion of hens recorded feather pecking in the 180min observation period was 75 versus 49% (P<0.001).In generation 3, plumage condition was better in LP on neck, breast, back, wings and tail, as well as overall (P<0.001). Body weight did not differ between lines in generation 2, but in generation 3, HP hens were on average heavier than LP hens at the age of 27 weeks (1435g versus 1371g, P<0.001).     

Lergotrile and lisuride: in vivo dopaminergic agonists which do not stimulate the presynaptic dopamine autoreceptor.

The presynaptic dopaminergic autoreceptor which regulates tyrosine hydroxylase is unaffected by the dopaminergic ergots and their specific antagonists. Apomorphine inhibits the conversion of tyrosine to catechols by striatal synaptosomes; neither lergotrile nor lisuride mimic this action of apomorphine. Furthermore metoclopramide, a potent dopaminergic antagonist in the anterior pituitary, does not block the inhibitory effect of apomorphine. The data demonstrate that the several categories pharmacologically distinct dopamine receptors exist.