Quantification of the contact structure in a feral cattle population and its hypothetical effect on the transmission of bovine herpesvirus 1.

The organisation of animal populations in social groupings may play a crucial role in the transmission of any infectious disease that requires close contact. The objective of this study was to quantify the contact structure of part of the Heck cattle population in a Dutch nature reserve and its hypothetical effect on the transmission of bovine herpesvirus 1 (BHV1). The contact structure was quantified by observing the number of different animals with whom contact was made (i.e. the number of contactees) within a fixed time period. Two types of behaviour sampling methods, namely focal sampling and scan sampling were used to observe the contact structure. In this study only those contacts between individuals were observed that were assumed to be a proxy measure of an at-risk event for BHV1-infection. Two reproduction ratios (R), i.e. the average number of new cases caused by a typical infected individual, were estimated, one for the observed contact structure and another for a random mixing contact structure. The two reproduction ratios were then compared to study the hypothetical effect on BHV1 transmission. The overall number of contactees was highest during summer and lowest during winter-spring. The contact structure of the homogeneous population did differ significantly from a random mixing contact structure, resulting in that the variation in the number of contactees was higher than under random mixing. Bulls, young bulls and cows had the highest number of contactees during, respectively, summer, autumn and winter-spring. From the analysis of the contingency tables it was clear that contacts between animal types did not occur at random during summer and autumn. For example, during summer more contacts than expected occurred between bulls and cows. This heterogeneity at animal type level was taken into account in the calculation for R, which resulted for the observed contact structure in higher estimates for R than for the homogeneous population. When looking at heterogeneity at individual level it was found that during summer almost all individuals were observed together direct or indirect in the same group except for certain bull groups. During autumn and winter-spring almost all individuals were seen together in the same group when considering a long contact period of 14 days but the groups were fallen apart in smaller groups and solitary individuals for a short contact period of 5 days. It could be concluded that based on the observed contact structure transmission would be favoured most during summer.

Temporal and sequential structure of behavior and facility usage of laying hens in an enriched environment.

Improved housing for laying hens may start from the translation of their behavioral needs into welfare-based design parameters for laying hen houses. The objective of our research was to gain insights into the facility usage and behavioral needs of the hen over 24 h when there are no obvious restraints. Twenty ISA Brown commercial laying hens (Gallus domesticus) that were 18 wk old and not beak trimmed, were accommodated in a pen (4 x 6 m) at 19 + 2 degrees C on a light-dark cycle of 10L:14D. The pen providing nest boxes, drinkers, feeders, perches, sand, and wood shaving was designed to accommodate the hens for the experimental period. Video recordings were made for 10 d. Behavioral analyses were conducted on 5 birds for 5 d. Time spent on each behavior, log survivor analysis of events and inter-event intervals, bout analysis, diurnal pattern in events and bouts, occurrence of behavior in different segments and the corridor of the pen, and sequence analysis were performed to gain insights into the temporal and sequential structures of behavior. Hens spent 97% of the day on nest use, preening, drinking, feeding, still, walking, perching, and resting; 43% on commodity-dependent behavior; and 57% not on commodity-oriented behaviors. Behavioral events were short (around 70% event <2 min) and frequent (around 70% inter-event intervals <40s). The pen corridor was the preferred place for attack, escape, flying, resting, walking, and wing flapping. Feeding-drinking-feeding, preening-resting-preening, scratching-resting-scratching, dust bathing-resting-preening, or dust bathing-resting-wing stretching-dust bathing were the preferred sequences of behavior. Although hens interrupted ongoing behaviors and changed behaviors frequently, they nonetheless clustered behavioral events.

Involvement of the nucleus accumbens in oral behaviour in the freely moving rat.

The role of the nucleus accumbens in oral behaviour was examined by intra-accumbens injections of a single dose of a selective dopamine D1 receptor agonist (SKF 38393: 5 micrograms/side), a selective dopamine D2 receptor agonist (quinpirole: 10 micrograms/side), and their combination in freely moving rats. Principal factor analysis revealed four factors to be involved in the scored behaviours, two of which concerned oral behaviour: a chew factor, comprising the behaviours chew, tongue protrusion, yawn and lick, and a groom factor, with high factor loadings of tremor and groom. The two remaining factors were the circle factor comprising circle, walk and rear, and the sniff factor comprising sniff, yawn and rear. Two-way ANOVA (independent variable D1 with H2O and SKF 38393 level; independent variable D2 with H2O and quinpirole level) of the factor scores revealed that SKF 38393 and quinpirole had similar or opposite effects which were additive or antagonistic, depending on which behaviour was studied. This study demonstrates that (a) the nucleus accumbens plays a major role in the oral behaviour of freely moving rats, and (b) an integrated study of all oral behavioural elements is necessary to describe the effects of drugs on oral behaviour.

Jaw muscle activity, the nucleus accumbens, and dopaminergic agonists: a new approach.

A method was developed to analyze electromyographic (EMG) signals in terms of power, viz., a measure for overall muscle activity, and number of seconds marked by distinct frequency ranges. With the help of this method, the effects of intraaccumbens administration of distilled water, the D1 receptor agonist SK&F 38393 (SKF; 5 micrograms), the D2 receptor agonist LY 171555 (LY; 10 micrograms), and their combination upon the EMG signals of the masseter and the digastric muscle were analysed in freely moving rats. Only the combined treatment affected the power: The noted increase was limited to the digastric muscle. The time/frequency analysis was limited to frequency ranges 3-4 Hz (class A), 4-5 and 5-6 Hz (class B), and 6-7, 7-8, ..., 12-13, and 13-14 Hz (class C). Apart from a small effect of SKF alone and of SKF in combination with LY on class B of the masseter muscle, neither SKF nor LY affected class A or B. SKF and LY increased and decreased, respectively, class C in both muscles. The data suggest that SKF and LY elicited both opposite and synergistic effects. The method is a new tool to analyze EMG signals in freely moving rats.

Temporal structure of red jungle fowl crow sequences: single-case analysis.

Vocalisations can be indicators of the physical state or condition of an animal, for instance the crow of a rooster. There are no published data on the temporal structure of crow sequences. In this paper 536 crows in 22 sequences of one red jungle fowl rooster (rooster 1) were recorded and analysed. Calls were very characteristic and there was little variation between the calls. Rooster 1 crowed with a mean latency of 287 s after onset of a light in the early morning. The average crow sequence consisted of 29 calls which lasted for 1416 ms on average. A gradual increase in pre-call interval was characteristic of each sequence. The variation in note durations is systematic and especially the last note correlated highly with the total call duration. Both the pre-call interval and the duration of the 4th note (doooo) may be parameters of fatigue and thus of physical condition. The number of crows in a sequence was positively related to air pressure outdoors. An experiment is done in which the physical condition of a rooster (rooster 2) is manipulated by food deprivation. The results are analysed with single case randomisation tests. The number of crows in a sequence is significantly reduced after food deprivation, while the pre-call intervals are significantly increased. Prediction of the number of crows in the sequence can be done on base of the pre-call interval of the second call: both rooster 1 and 2 showed long pre-call intervals in short call sequences. Food deprivation and the possible loss of condition have a significant effect on the temporal structure of crow sequences and cause a decrease in the number of crows in a call sequence.

The effect of frustrative nonreward on vocalisations and behaviour in the laying hen, Gallus gallus domesticus.

Laying hens are thought to express an expectation of a rewarding event through a specific vocalisation, the gakel-call. It has been suggested that the gakel-call is related to frustration, i.e. the thwarting of behaviour. We investigated if frustrative nonreward (nonreinforcement in a situation that previously was consistently reinforcing) in laying hens is expressed through this gakel-call. Twenty hens of two commercial strains, ten ISA White Leghorn and ten ISA Brown Warren were subjected to a classical conditioning procedure. After 23 h of food-deprivation they were trained, in automated Skinnerboxes, to use red lights as a signal (unconditioned stimulus) for a food reward. After this the hens of each strain were equally divided into two groups of five. They were subjected to a control session (light as conditioned stimulus followed by food reward) and to a frustration session (conditioned stimulus followed by nonreward). Both during the frustration and control session behaviour and vocalisations were recorded. In the White Leghorn strain high levels of alarm-calls are found as indicators of anxiety. In the Brown Warren strain more gakel-calls and an increased locomotor activity are found after frustrative nonreward. In conclusion, both findings suggest that vocalisations could serve as indicators of a laying hen's welfare.

Reaction to frustration in high and low feather pecking lines of laying hens from commercial or semi-natural rearing conditions.

The effect of rearing conditions on feather pecking and reaction to frustration was studied in two lines of laying hens. From commercial rearing conditions (large group, no mother hen), seven birds from a high feather pecking line (HC birds) and eight birds from a low feather pecking line (LC birds) were used. From semi-natural rearing conditions (small group, mother hen present) seven birds from the high feather pecking line (HN birds) were used. Feather pecking behaviour of HC, LC, and HN groups was recorded for 30 min. After that, each bird was food deprived and trained to peck a key for a food reward in a Skinnerbox. After training, each bird was subjected to a frustration session in a Skinnerbox, where the feeder was covered with Perspex. Three HC birds showed severe feather pecking, compared with one HN bird and zero LC birds. Differences in reaction to frustration were found between birds from different lines, but not in birds from different rearing conditions. LC birds tended to put their head in the feeder more frequently than HC birds over all sessions. Although limited, this study indicates that rearing conditions influence feather pecking, but not reaction to frustration.

Heritability of feather pecking and open-field response of laying hens at two different ages.

The objective of the current study was to estimate heritabilities (h2) of feather pecking and open-field response of laying hens at two different ages. An F2 cross, originating from a high and a low feather pecking line of laying hens, was used for the experiment. Each of the 630 birds of the F2 cross was subjected to an open-field test (individual, 10 min) at 5 and 29 wk of age and to a social feather pecking test (groups of five birds on wood shavings, 30 min) at 6 and 30 wk of age. Both tests were performed in a square open field (1.25 x 1.25 m). Behavior was recorded directly from a monitor. Heritabilities of feather pecking and open-field behaviors were calculated. In the open-field test at 5 wk of age, high h2 were found for most traits, ranging from 0.20 for the frequency of flying to 0.49 for number of steps. In the social test at 6 wk, gentle feather pecking (0.12) and ground pecking (0.13) were found to be heritable. When both tests were repeated at 29 and 30 wk of age, h2 estimates were lower for the open-field test, ranging from 0.10 for duration of sitting to 0.20 for latency to first step. In the social test, however, higher h2 estimates of 0.15 for gentle feather pecking and 0.30 for ground pecking were found compared with 6 wk of age. In conclusion, gentle feather pecking and open-field behaviors may be used in selection against feather pecking.

Genetic and phenotypic correlations between feather pecking behavior, stress response, immune response, and egg quality traits in laying hens.

The objective of the current study was to estimate genetic and phenotypic correlations among feather pecking (FP) behavior and stress response, immune response, and egg quality parameters. These traits have been measured in an F2 cross, coming from a cross between a high and a low FP line of laying hens. Heritabilities (h2) of stress response (32 wk), primary immune response to keyhole limpet hemocyanin (KLH) (36 wk) and Mycobacterium butyricum (39 wk), and egg quality parameters (35, 44, and 50 wk of age) were calculated. The h2 was 0.05 +/- 0.05 (SE) for stress response, 0.15 +/- 0.07 for antibody response to KLH, and 0.08 +/- 0.06 for antibody response to M. butyricum. The h2 for egg quality traits were in the range of 0.12 to 0.30. Significant phenotypic correlations were found between gentle FP in adult hens and stress response, egg weight at 44 and 50 wk, and egg deformation at 50 wk. Significant additive genetic correlations were found between severe FP in adult hens and antibody response to KLH (0.79 +/- 0.35), and between ground pecking in adult hens and egg deformation at 50 wk (0.63 +/- 0.26), and between ground pecking and eggshell strength at 35, 44, and 50 wk of age (-0.86 +/- 0.29, -0.81 +/- 0.20, -0.76 +/- 0.24, respectively).

Identification of quantitative trait loci for receiving pecks in young and adult laying hens.

Feather pecking (FP) is a major problem in cage and free-range housing systems. In free-range systems, FP is more difficult to control. It is not known why a victim is being pecked. It could be that a bird is genetically predisposed to be pecked. To study the genetics of FP behavior, a large F2 population of 630 hens was generated from a cross between two commercial laying lines differing in their propensity to feather peck. The traits measured at 6 and 30 wk of age were receiving gentle FP, receiving severe FP, and receiving aggressive pecking. In addition, receiving toe pecking (TP) was also measured at 30 wk of age. For receiving gentle FP at 6 wk of age, a significant QTL on GGA1 and three different suggestive QTL were identified on GGA2, GGA6, and GGA7, respectively. For receiving gentle FP at 30 wk of age, a suggestive QTL on GGA5 was detected. For receiving aggressive pecking, a suggestive QTL was detected on GGA3. For receiving TP, three suggestive QTL were detected on GGA1, GGA5, and GGA23, respectively. The different QTL detected for receiving gentle FP at different ages indicate that this trait is regulated by different genes in young and adult hens.

Mapping quantitative trait loci affecting feather pecking behavior and stress response in laying hens.

In the European Union, legislation concerning animal housing is becoming stricter because of animal welfare concerns. Feather pecking (FP) in large group housing systems is a major problem. It has been suggested that corticosterone (CORT) response to manual restraint as a measure for stress is associated with FP behavior. The aim of the current study was to identify QTL involved in FP behavior and stress response in laying hens. An F2 population of 630 hens was established from a cross between two commercial lines of laying hens differing in their propensity to feather peck. The behavioral traits, measured at 6 and 30 wk of age, were gentle FP, severe FP, and aggressive pecking. Toe pecking was measured at 30 wk of age and CORT response to manual restraint was measured at 32 wk. All animals were genotyped for 180 microsatellite markers. A QTL analysis was performed using a regression interval mapping method. At 6 wk of age, a suggestive QTL on GGA10 was detected for gentle FP. At 30 wk of age, suggestive QTL were detected on GGA1 and GGA2 for gentle FP. A significant QTL was detected on GGA2 for severe FP. At 32 wk of age, a suggestive QTL was detected on GGA18 for CORT response to manual restraint. In addition, a suggestive QTL was detected on GGA5 with possible maternal parent-of-origin effect for CORT response.

Thwarting of behaviour in different contexts and the gakel-call in the laying hen.

Earlier studies have shown that thwarting of feeding behaviour in the laying hen is expressed through a specific vocalisation, the gakel-call. The first aim of this study was to investigate whether the effect of deprivation per se on the occurrence of gakel-calls can be distinguished from the effect of the additional frustration. Frustration is defined as the state of an animal that results from nonreward in the expectancy of reward. The second aim was to investigate whether the occurrence of gakel-calls is restricted to a food context or whether it can be regarded as an expression of frustration in general. For this purpose, 20 hens were deprived of food, water and dustbath. After deprivation at a fixed time, a cue was given and the hens were rewarded with access to food, water or dust during a 15-min session on 4 consecutive days. On the fifth day, they were thwarted in the associated behaviours by blocking the access to these commodities, after the hens had been presented the signal that previously preceded the reward. We then recorded behaviours that might reflect the state of frustration in three 15-min periods. The period "Pre-Frustration" started 15 min before "Frustration". This, in turn, was followed by the period "Post-frustration" in which the hens were rewarded again. Nesting behaviour was thwarted by blocking the access to the nest (Frustration) after a hen had reached the last stage of its prelaying behaviour.In the food, water and dustbath context, deprivation elicited gakel-calls. The additional frustration resulted in a higher number of gakel-calls in all contexts except the food context. However, together with the findings of previous experiments, the results of this study suggest that frustration, in general, is expressed through the gakel-call. Frustration in the nest context elicited more gakel-calls than the other contexts. This latter finding is discussed in the light of the occurrence of the gakel-call under natural circumstances.

The vocal expression of feeding motivation and frustration in the domestic laying hen, Gallus gallus domesticus.

Thwarting of feeding behaviour in the laying hen results in an increase in stereotyped pacing, displacement preening, and the gakel-call. These behaviours therefore reflect the frustration arousal caused by the thwarting of feeding behaviour. This raises the question whether the level of frustration also varies with the intensity of the motivation to perform the thwarted behaviour. This study investigated the relationship between the intensity of the motivation and level of frustration on the one hand and the gakel-call on the other hand. In Experiment 1, the strength of the motivation to feed was varied by thwarting hens in their feeding behaviour in an operant procedure after different durations of food deprivation (0, 8, 23 and 47 h). Trend analysis showed that with increasing hunger state, an increasing number of gakel-calls was given. No effect of treatments on temporal characteristics of the gakel-call was found. In Experiment 2, the level of frustration was varied by reducing or increasing the duration of access to food for food-deprived hens compared to the duration of access during training. It was assumed that a shorter duration of access to food compared to training would elicit frustration, which in turn would affect the performance of behaviours indicative of thwarting. However, we found neither a relation between the number of gakel-calls nor the temporal features of the gakel-call and the duration of access to food. Possibly, the differences between treatments were not large enough to induce differences in frustration level. Also, other factors that might have influenced the motivation are discussed.

Eating behaviour, and preprandial and postprandial correlations in male broiler and layer chickens.

1. It has been suggested that broiler chickens have a disturbed satiety and hunger mechanism. The satiety mechanism for eating can be expressed as the positive correlation between meal length and the length of the preceding (preprandial) interval; the hunger mechanism for eating as the positive correlation between meal length and the length of the succeeding (postprandial) interval. An experiment was conducted to investigate eating behaviour of male broiler and layer chickens by measuring meal and interval lengths. 2. Eight male broilers and 8 male layer chickens were housed individually and visually isolated in floor pens (1 m2/pen) on wood shavings. From 4 to 7 weeks of age, eating behaviour of each bird was recorded for 3 h in two conditions each week. In the first condition, the birds were not deprived from feed. In the second condition, they were 24-h food deprived and feed was provided just before the observation started. Preprandial and postprandial correlations were calculated based on data of the non-deprived condition. Before and after each observation bird and feeder were weighed to measure weight gain and feed consumption during observation. 3. Under the non-deprived condition, the broilers spent initially more, but at a later age less time on eating. The broilers had fewer meals per hour, consumed more feed per hour, and had longer meal and interval lengths than the layer chickens. After 24-h feed deprivation, the broilers had a longer first meal, consumed more feed per hour and spent more time on eating than the layer chickens. Significant preprandial correlations but no postprandial correlations were found in the broilers. In the layer chickens, both significant preprandial and postprandial correlations were found. This indicates that for regulating eating behaviour, the satiety mechanism dominates the hunger mechanism in broilers, and satiety and hunger mechanisms are equally involved in layer chickens. 4. The typical eating behaviour of broilers and the calculated preprandial and postprandial correlations have given new indications that hunger and satiety mechanisms in broilers have changed compared with layer chickens. In broilers, there is no lower set point, but only an upper set point for controlling eating behaviour, which suggests that broilers eat to their maximal physical capacity.

Strain differences in rats with respect to speed of conflict resolution.

Speed of conflict resolution was studied in a conditioned punishment paradigm in a Skinner box and a straight runway. In both experimental situations speed of conflict resolution was defined as the latency to gain food during an approach-avoidance conflict. In the Skinner box Tryon Maze Bright rats were faster in speed of conflict resolution than Tryon Maze Dull rats, and Roman Low Avoidance rats were faster than Roman High Avoidance rats. In the runway situation, Wistar Kyoto rats were faster in solving the conflict than randomly bred Wistar Wu rats and Brown Norway rats were faster than Wistar Wu rats. Differences between the strains in speed of conflict resolution could not be consistently explained from strain differences in approach or avoidance behavior, measured separately. It is, therefore, suggested that speed of conflict resolution is a unique parameter.

Reaction to frustration in high and low feather pecking laying hens.

Reaction to frustration of high (HFP) and low feather pecking (LFP) laying hens was investigated. From a HFP- and a LFP-line five birds with a HFP- and five birds with a LFP-phenotype were selected. Birds from the HFP-line were expected to show more key pecking and covered feeder pecking during frustration than birds from the LFP-line. When a bunch of feathers was presented, birds with a HFP-phenotype were expected to redirect their pecks at the bunch. Birds were trained to peck a key for a food reward in an automated Skinnerbox and subjected to two sessions: a control session, where food was available, and a frustration session, where the feeder was covered with Perspex. These two sessions were repeated in the presence of a bunch of feathers. Unexpectedly, birds from the LFP-line had a stronger reaction to frustration than birds from the HFP-line, expressed in pecking behaviour. When a bunch of feathers was offered, birds with a HFP-phenotype did not show more bunch pecking during frustration than birds with a LFP-phenotype.

Genetic and phenotypic correlations between feather pecking and open-field response in laying hens at two different ages.

The object of this research was to study the relationship between feather pecking and open-field activity in laying hens at two different ages. A population of 550 birds of a laying hen cross was subjected to an open-field test at 5 and 29 weeks of age and to a social feather pecking test at 6 and 30 weeks of age. Factor analysis was used to identify underlying factors for each test: pecking behavior (social test) and open-field activity (open-field test). In young birds, a positive phenotypic correlation of 0.24 was found between high open-field activity and high levels of pecking behavior (ground pecking, preening, gentle feather pecking, and wall pecking). In adults, a similar genetic correlation of 0.62 was found. At adult age, the factor pecking behavior consisted mainly of gentle and severe feather pecking. Between ages, a strong, negative genetic correlation of -0.65 was found between open-field activity at young age and pecking behavior at adult age, indicating that open-field activity levels in young birds may predict pecking behavior in adult hens.

Identification of QTLs involved in open-field behavior in young and adult laying hens.

Line differences for open-field behavior in chickens have been observed, and it has been shown that this behavior has a genetic component. The aim of this study was to detect quantitative trait loci (QTL) involved in open-field behavior. For this purpose, open-field behavior was studied at 5 and 29 weeks of age in F(2) hens coming from an intercross between two commercial White Leghorn laying lines selected for egg production traits. Latencies, durations, and frequencies of general activity (sitting, standing, walking, and stepping), defecation, and vocalizations were recorded individually for each bird, and a factor score was calculated. All animals (F(0), F(1), and F(2)) were screened with 180 microsatellite markers. Regression interval mapping was applied using both a paternal half-sib analysis and a line-cross analysis method. For general activity at 5 weeks of age, a significant QTL was detected on GGA4 and a suggestive QTL on GGA2 under the line-cross model. For general activity at 29 weeks of age, a significant QTL was detected on GGA4 and two suggestive QTLs were detected on GGA1 and on GGA10, respectively, also using the line-cross analysis. The QTL on GGA4 at 5 weeks of age did not overlap with the QTL on GGA4 at 29 weeks of age. The current study indicated that open-field behavior in young chickens was regulated by QTL that differ from the QTL for open-field behavior in adult chickens.