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Research article2024Peer reviewedOpen access

Comb size, shape complexity and laterality of laying hens reared in environments varying in resource choice

Holt, R. V.; Skanberg, L.; Keeling, L. J.; Estevez, I.; Lepej, P.; Andersen, I. L.; Vas, J.; Newberry, R. C.


The comb is an ornament involved in signalling condition in domestic fowl. We hypothesised that comb size, comb shape complexity (i.e., rugosity, the comb perimeter jaggedness), and comb laterality of laying hens would be influenced by the degree of environmental enrichment experienced during juvenile development in the form of resource choice. We conducted a 2 x 2 factorial crossover experiment with pullets reared in pens containing four perches of equal length and four litter areas of equal size. Pullets were exposed to a single choice vs multiple choices of perch and litter types (i.e., all the same vs all different) during Weeks 1-4 (Period 1) and/or Weeks 5-15 (Period 2) of rearing (n = 4 pens/treatment combination) prior to transfer to standard adult laying pens for Weeks 16-27 (Period 3). In Week 27, combs were photographed, and comb laterality (hanging on left or right side) was noted. Using a custom-made image analysis programme, we captured comb area (mm 2 ), perimeter length (mm), and rugosity ((perimeter length - horizontal length) / horizontal length) from comb photographs of 6-7 randomly selected hens/pen. We predicted that hens reared in the multi -choice environment during Periods 1 and 2 would have larger, more complex, and left -side -biased combs than those in the other treatment groups, reflecting lower allostatic load. The predicted comb side bias was based on a possible bias in head posture/movements associated with greater right eye/ear use and left -brain hemispheric dominance. Contrary to our predictions, we detected an overall right -side bias in comb laterality, and no associations between resource choice treatment in Period 1 or Period 2 and comb area, perimeter length, rugosity, or laterality of the adult hens. Thus, variation in allostatic load resulting from the rearing treatments was insufficient to modify the trajectory of comb morphological development, possibly due to a ceiling effect when comparing environmental treatments on the positive end of the welfare spectrum. We found that left -lopping combs had shorter perimeters than right -lopping combs. However, among hens with left -lopping combs, those with larger combs were heavier and had less feather damage, while among hens with right -lopping combs, those with longer -perimeter combs were heavier and tended to have less comb damage. In conclusion, comb characteristics were related to physical condition at the individual level but did not serve as sensitive integrated indicators of hen welfare in response to basic vs enhanced resource choice during rearing. (c) 2024 The Author(s). Published by Elsevier B.V. on behalf of The Animal Consortium. This is an open access article under the CC BY license (


Condition-dependent signalling; Environmental enrichment; Female ornamentation; Positive animal welfare; Stress resilience

Published in

2024, Volume: 18, number: 6, article number: 101157
Publisher: ELSEVIER