Karin E. Schütz

Domestication effects on foraging strategy, social behaviour and different fear responses: a comparison between the red junglefowl (Gallus gallus) and a modern layer strain

Abstract According to the resource allocation theory, behaviours that have high energetic costs should decrease in frequency in breeds that are selected to invest a higher proportion of energy into production traits. We studied differences between red junglefowl, ancestor of domesticated breeds, and a modern White Leghorn laying strain with respect to foraging strategy, social behaviour and different fear responses. The behaviour of each breed was observed in four test situations: (1) a foraging-social maze of four arms where two arms contained easily obtainable familiar food and two contained hidden novel food, so the birds had to work to obtain it. One arm with each food type had a mirror, which provided a simulated social stimulus. (2) A hawk model test, where a hawk model was briefly presented while the birds had access to food. (3) A tonic immobility test, measuring passive fear responses. (4) A restraint test whereby the birds were restrained with a rope around one tarsus, measuring active fear responses. In the maze test, junglefowl spent more time feeding from the novel food site and next to a mirror, whereas the Leghorn spent more time in the arm with the easily obtainable familiar food and mirror. Both breeds responded on the simulated predator attack with an immediate decrease in foraging behaviour and an increase in “walking alert”. After the model presentation, junglefowl walked alert and vocalised significantly more than the Leghorn, whereas Leghorn was standing alert more compared to junglefowl. It was easier to induce tonic immobility in the Leghorn than in junglefowl and Leghorn also had a less active response in the restraint test compared to junglefowl. Females in general tended to respond more active than males. The results on social behaviour were not clear. In conclusion, Leghorn used a more energy-conserving foraging strategy and was less active in the fear tests, compared to junglefowl. These results may be interpreted as correlated responses to selection for increased production, allowing modern strains to allocate more resources to reproduction and growth.

Chicken genomics: Feather-pecking and victim pigmentation

Feather-pecking in domestic birds is associated with cannibalism and severe welfare problems. It is a dramatic example of a spiteful behaviour in which the victims fitness is reduced for no immediate direct benefit to the perpetrator and its evolution is unexplained. Here we show that the plumage pigmentation of a chicken may predispose it to become a victim: birds suffer more drastic feather-pecking when the colour of their plumage is due to the expression of a wild recessive allele at PMEL17, a gene that controls plumage melanization, and when these birds are relatively common in a flock. These findings, obtained using an intercross between a domestic fowl and its wild ancestor, have implications for the welfare of domestic species and offer insight into the genetic changes associated with the evolution of feather-pecking during the early stages of domestication.

Feather pecking in chickens is genetically related to behavioural and developmental traits

Feather pecking (FP) is a detrimental behaviour in chickens, which is performed by only some individuals in a flock. FP was studied in 54 red junglefowl (ancestor of domestic chickens), 36 White Leghorn laying hens, and 762 birds from an F(2)-intercross between these two lines. From all F(2)-birds, growth and feed consumption were measured. Age at sexual maturity and egg production in females, and corticosterone levels in males were also measured. From 333 F(2)-birds of both sexes, and 20 parental birds, body composition with respect to bone mineral content, muscle and fat was obtained by post-mortem examinations using Dual X-Ray Absorptiometry (DXA). In femurs of the same birds, the bone density and structure were analysed using DXA and Peripheral Quantitative Computerized Tomography (pQCT), and a biomechanical analysis of bone strength was performed. Furthermore, plumage condition was determined in all birds as a measure of being exposed to feather pecking. Using 105 DNA-markers in all F(2)-birds, a genome-wide scan for Quantitative Trait Loci (QTL), associated with the behaviour in the F(2)-generation was performed. FP was at least as frequent in the red junglefowl as in the White Leghorn strain studied here, and significantly more common among females both in the parental strains and in the F(2)-generation. In the F(2)-birds, FP was phenotypically linked to early sexual maturation, fast growth, weak bones, and, in males, also high fat accumulation, indicating that feather peckers have a different resource allocation pattern. Behaviourally, F(2) feather peckers were more active in an open field test, in a novel food/novel object test, and in a restraint test, indicating that feather pecking might be genetically linked to a proactive coping strategy. Only one suggestive QTL with a low explanatory value was found on chromosome 3, showing that many genes, each with a small effect, are probably involved in the causation of feather pecking. There were significant effects of sire and dam on the risk of being a victim of feather pecking, and victims grew faster pre- and post-hatching, had lower corticosterone levels and were less active in a restraint test. Hence, a wide array of behavioural and developmental traits were genetically linked to FP.

The amount of shade influences the behavior and physiology of dairy cattle

The objective was to understand how the amount of shade (shade cloth blocking 99% of solar radiation) influenced the behavior and physiology of Holstein-Friesian dairy cattle managed on pasture. We compared behavior, body temperature, and respiration rate of cattle provided with 1 of 3 treatments for 5 d: access to 2.4m(2) or 9.6m(2) shade/cow, or no shade (n=4 groups/treatment, 10 animals/group). Behavioral observations were carried out between 1000 and 1550h. Cows spent more than twice as much time in the larger shade (24 vs. 50% of observations for 2.4m(2) and 9.6m(2) shade/cow, respectively, SED: 1.7%) and engaged in fewer aggressive interactions when more shade was provided (10.7 vs. 3.2 aggressive interactions/m(2) during 5.8h of observation for 2.4m(2) and 9.6m(2) shade/cow, respectively, SED: 3.16 interactions/m(2)). Time around the water trough increased when little or no shade was provided (11, 5, and 2% of observations within 4.5m of water trough for no shade, 2.4m(2), and 9.6m(2) shade/cow, SED: 2.4%). Respiration rate was higher when cows had less shade available (62, 57, and 51 breaths/min for no shade, 2.4m(2), and 9.6m(2) shade/cow, respectively, SED: 2.1 breaths/min). All cows used the shade more when 9.6m(2) shade/cow was provided; simultaneous use was observed in 15 versus 0% of observations in the 9.6m(2) and 2.4m(2) treatments on the warmest day, respectively. Weather conditions influenced both the behavioral and physiological responses, and these changes were more pronounced when less or no shade was available. Cows spent more time in shade and less time lying with increasing heat load. In addition, aggressive interactions in the shade, time around the water trough, mean body temperature, and respiration rate increased with environmental heat load. Our findings highlight the importance of determining and providing an effective amount of shade to cattle.

The genetic architecture of a female sexual ornament

Abstract Understanding the evolution of sexual ornaments, and particularly that of female sexual ornaments, is an enduring challenge in evolutionary biology. Key to this challenge are establishing the relationship between ornament expression and female reproductive investment, and determining the genetic basis underpinning such relationship. Advances in genomics provide unprecedented opportunities to study the genetic architecture of sexual ornaments in model species. Here, we present a quantitative trait locus (QTL) analysis of a female sexual ornament, the comb of the fowl, Gallus gallus, using a large-scale intercross between red junglefowl and a domestic line, selected for egg production. First, we demonstrate that female somatic investment in comb reflects female reproductive investment. Despite a trade-off between reproductive and skeletal investment mediated by the mobilization of skeletal minerals for egg production, females with proportionally large combs also had relatively high skeletal investment. Second, we identify a major QTL for bisexual expression of comb mass and several QTL specific to female comb mass. Importantly, QTL for comb mass were nonrandomly clustered with QTL for female reproductive and skeletal investment on chromosomes one and three. Together, these results shed light onto the physiological and genetic architecture of a female ornament.

Dairy cattle prefer shade over sprinklers: Effects on behavior and physiology

Cattle will readily use shade in warm weather, but less is known about voluntary use of sprinklers. We examined preferences of 96 Holstein-Friesian dairy cows (milk yield: 12.7±3.48 kg per day; mean±SD) for sprinklers, shade, or ambient conditions after walking 2.0 km or 0.3 km before afternoon milking (n=48 cows/distance). Each cow was individually tested on 3 consecutive days with a different paired choice each day: 1) shade or sprinklers, 2) shade or ambient conditions, 3) sprinklers or ambient conditions. Average air temperature during testing was 22.3°C. Cows preferred shade over sprinklers (62 vs. 38% ± 5.0%; mean ± SE) and shade over ambient conditions (65 vs. 35% ± 5.1%; mean±SE). Cows showed no preference between sprinklers and ambient conditions (44% of the cows chose sprinklers, SE=5.3%). The preference for shade over sprinklers and ambient conditions increased with air temperature, solar radiation, and wind speed. Walking distance did not influence the preference for any treatment. Respiration rate was decreased most by sprinklers (38% decrease) but also decreased in shade and ambient conditions (17 and 13% decrease, respectively; standard error of the difference=4.7%). Similarly, surface temperature was decreased most by sprinklers (11.4% decrease), compared with that by shade (1.0% decrease), or that by ambient conditions (1.4% increase; standard error of the difference=1.82%). Furthermore, sprinklers reduced insect avoidance behaviors, including number of tail flicks and hoof stamps. In conclusion, dairy cattle preferred to use shade in summer despite sprinklers being more efficient in decreasing heat load and insect avoidance behavior.

The genetic architecture of domestication in the chicken: effects of pleiotropy and linkage

The extent of pleiotropy and epistasis in quantitative traits remains equivocal. In the case of pleiotropy, multiple quantitative trait loci are often taken to be pleiotropic if their confidence intervals overlap, without formal statistical tests being used to ascertain if these overlapping loci are statistically significantly pleiotropic. Additionally, the degree to which the genetic correlations between phenotypic traits are reflected in these pleiotropic quantitative trait loci is often variable, especially in the case of antagonistic pleiotropy. Similarly, the extent of epistasis in various morphological, behavioural and life‐history traits is also debated, with a general problem being the sample sizes required to detect such effects. Domestication involves a large number of trade‐offs, which are reflected in numerous behavioural, morphological and life‐history traits which have evolved as a consequence of adaptation to selective pressures exerted by humans and captivity. The comparison between wild and domestic animals allows the genetic analysis of the traits that differ between these population types, as well as being a general model of evolution. Using a large F2 intercross between wild and domesticated chickens, in combination with a dense SNP and microsatellite marker map, both pleiotropy and epistasis were analysed. The majority of traits were found to segregate in 11 tight ‘blocks’ and reflected the trade‐offs associated with domestication. These blocks were shown to have a pleiotropic ‘core’ surrounded by more loosely linked loci. In contrast, epistatic interactions were almost entirely absent, with only six pairs identified over all traits analysed. These results give insights both into the extent of such blocks in evolution and the development of domestication itself.

Quantitative Trait Loci for BMD and Bone Strength in an Intercross Between Domestic and Wildtype Chickens

With chicken used as a model species, we used QTL analysis to examine the genetic contribution to bone traits. We report the identification of four QTLs for femoral traits: one for bone strength, one for endosteal circumference, and two affecting mineral density of noncortical bone.

Body condition score at calving affects systemic and hepatic transcriptome indicators of inflammation and nutrient metabolism in grazing dairy cows.

Calving body condition score (BCS) is an important determinant of early-lactation dry matter intake, milk yield, and disease incidence. The current study investigated the metabolic and molecular changes induced by the change in BCS. A group of cows of mixed age and breed were managed from the second half of the previous lactation to achieve mean group BCS (10-point scale) that were high (HBCS, 5.5; n=20), medium (MBCS, 4.5; n=18), or low (LBCS, 3.5; n=19). Blood was sampled at wk -4, -3, -2, 1, 3, 5, and 6 relative to parturition to measure biomarkers of energy balance, inflammation, and liver function. Liver was biopsied on wk 1, 3, and 5 relative to parturition, and 10 cows per BCS group were used for transcript profiling via quantitative PCR. Cows in HBCS and MBCS produced more milk and had greater concentrations of nonesterified fatty acids and β-hydroxybutyrate postpartum than LBCS. Peak concentrations of nonesterified fatty acids and β-hydroxybutyrate and greater hepatic triacylglycerol concentrations were recorded in HBCS at wk 3. Consistent with blood biomarkers, HBCS and MBCS had greater expression of genes associated with fatty acid oxidation (CPT1A, ACOX1), ketogenesis (HMGCS2), and hepatokines (FGF21, ANGPTL4), whereas HBCS had the lowest expression of APOB (lipoprotein transport). Greater expression during early lactation of BBOX1 in MBCS and LBCS suggested greater de novo carnitine synthesis. The greater BCS was associated with lower expression of growth hormone/insulin-like growth factor-1 signaling axis genes (GHR1A, IGF1, and IGFALS) and greater expression of gluconeogenic genes. These likely contributed to the higher milk production and greater gluconeogenesis. Despite greater serum haptoglobin around calving, cows in HBCS and MBCS had greater blood albumin. Cows in MBCS, however, had a higher albumin:globulin ratio, probably indicating a less pronounced inflammatory status and better liver function. The marked decrease in expression of NFKB1, STAT3, HP, and SAA3 coupled with the increase in ALB on wk 3 in MBCS cows were consistent with blood measures. Overall, results suggest that the greater milk production of cows with higher calving BCS is associated with a proinflammatory response without negatively affecting expression of genes related to metabolism and the growth hormone/insulin-like growth factor-1 axis. Results highlight the sensitivity of indicators of metabolic health and inflammatory state to subtle changes in calving BCS and, collectively, indicate a suboptimal health status in cows calving at either BCS 3.5 or 5.5 relative to BCS 4.5.

Calving body condition score affects indicators of health in grazing dairy cows

The objectives of this study were to determine the effect of calving body condition score (BCS) on cow health during the transition period in a pasture-based dairying system. Feed inputs were managed during the second half of the previous lactation so that BCS differed at drying off (BCS 5.0, 4.0, and 3.0 for high, medium, and low treatments, respectively: a 10-point scale); feed allowance was managed after cows were dried off, such that the BCS differences established during lactation remained at the subsequent calving (BCS 5.5, 4.5, and 3.5; n=20, 18, and 19, for high, medium, and low treatments, respectively). After calving, cows were allocated pasture and pasture silage to ensure grazing residuals >1,600 kg of DM/ha. Milk production was measured weekly; blood was sampled regularly pre- and postpartum to measure indicators of health, and udder and uterine health were evaluated during the 6 wk after calving. Milk weight, fat, protein, and lactose yields, and fat content increased with calving BCS during the first 6 wk of lactation. The effect of calving BCS on the metabolic profile was nonlinear. Before calving, cows in the low group had lower mean plasma β-hydroxybutyrate and serum Mg concentrations and greater mean serum urea than cows in the medium and high BCS groups, which did not differ from each other. During the 6 wk after calving, cows in the low group had lower serum albumin and fructosamine concentrations than cows in the other 2 treatment groups, whereas cows in the low- and medium-BCS groups had proportionately more polymorphonucleated cells in their uterine secretions at 3 and 5 wk postpartum than high-BCS cows. In comparison, plasma β-hydroxybutyrate and nonesterified fatty acid concentrations increased linearly in early lactation with calving BCS, consistent with a greater negative energy balance in these cows. Many of the parameters measured did not vary with BCS. The results highlight that calving BCS and, therefore, BCS through early lactation are not effective indicators of functional welfare, with the analyses presented indicating that both low and high BCS at calving will increase the risk of disease: cows in the low group were more prone to reproductive compromise and fatter cows had an increased risk of metabolic diseases. These results are important in defining the welfare consequences of cow BCS.