Nigel J. Ball

A Synthesis of Sleep in Wild Birds

In this synthesis we have attempted to survey and reanalyse the current literature concerning bird sleep. This was achieved by first reviewing present theory describing functions of sleep. Two general models surfaced which encompass many of the specific sleep theories, namely homeostatic and ecological theories. The terminology used to describe sleep and other associated behaviour was organized into an operational hierarchy. We defined loafing behaviour as a general state of immobility. This functional heterogenous group may include comfort behaviour, feeding, social interaction, wariness, staring down and inactive behaviour. Sleep and associated behaviour are considered inactive behaviour. We next described the common sleeping postures as follows: 1. Bill on back or under scapulars; the classical sleep posture. 2. Bill forward or rest-sleep posture. 3. Head on ground. Specific sleep characteristics were also identified as: 1. Quality of sleep based on duration and frequency of eyelid closure. 2. Length of sleep, called sleep session time which is based on the summation of sleep bouts. A sleep bout is defined as the time elapsed during eye closure and opening. 3. Temporal patterning of sleep sessions which are suggested to be dependent on timing of other higher priority behaviours. 4. The resilience (flexibility) of sleep session time. An original analysis was conducted on these sleep characteristics and other environmental, constitutional, behavioural and predation factors. The results indicated that birds experiencing longer daylength slept less each day. Further, birds slept less if (1) overall dangers were greater and (2) they were more communal sleepers. Longer sleep session times were associated with greater exposure while sleeping (i.e. on a lake or mud flat). Several possible functions of sleep were then suggested. Protection theories encompass choice of a safe sleep site while addressing the significance of communal roosting. Recuperation theories were not adequately supported by quantitative studies. Energy conservation theories imply that birds sleep when doing other behaviour might be non-productive. We suggest a model based on eyelid blinking which allows for a degree of vigilance during sleep but which is also compatible with minimizing energy expenditure.

Changing Heart Rates of Herring Gulls when Approached by Humans

Abstract Herring gulls (Larus argentatus) respond to an approaching human by an ‘emotional’ tachycardia. This response, which did not habituate, had a form dependent on the distance between the gull and an approaching human. It could also be modified by changing the speed of approach. Because there are reasons for believing this to be of adaptive significance, it is suggested that the response can be used in the study of animal behavior.

Manipulating daytime sleep in herring gulls (Larus argentatus)

Abstract The regular provision of a sufficient quantity of food on the territory of pairs of breeding herring gulls led to an increase in the time individuals spent on the territory during the day, relative to controls. This ‘extra’ time was not used to increase overall sleeping time, as predicted. Instead, the experimental gulls reduced their overall sleep time due to a decline in front-sleep. Back-sleep remained relatively unaffected, with both groups spending similar amounts of time in this posture. The consequences of the two postural types of sleep were examined with respect to the cost of allowing intruders to remain on an owners territory. Intruders arrived more often on control territories and stayed longer when only one pair member was present. Intruders were seen on control territories for more consecutive scans when the solitary control gull was in a sleep posture, particularly front-sleep. It is concluded that these results imply a difference in the consequences resulting from the two sleep postures and give additional support for the multiple function hypothesis of sleep.