title: Homeostatic regulation of sleep in the white-crowned sparrow (Zonotrichia leucophrys gambelii).
abstract: BACKGROUND
  Sleep is regulated by both a circadian and a homeostatic process. The homeostatic process reflects the duration of prior wakefulness: the longer one stays awake, the longer and/or more intense is subsequent sleep. In mammals, the best marker of the homeostatic sleep drive is slow wave activity (SWA), the electroencephalographic (EEG) power spectrum in the 0.5-4 Hz frequency range during non-rapid eye movement (NREM) sleep. In mammals, NREM sleep SWA is high at sleep onset, when sleep pressure is high, and decreases progressively to reach low levels in late sleep. Moreover, SWA increases further with sleep deprivation, when sleep also becomes less fragmented (the duration of sleep episodes increases, and the number of brief awakenings decreases). Although avian and mammalian sleep share several features, the evidence of a clear homeostatic response to sleep loss has been conflicting in the few avian species studied so far. The aim of the current study was therefore to ascertain whether established markers of sleep homeostasis in mammals are also present in the white-crowned sparrow (Zonotrichia leucophrys gambelii), a migratory songbird of the order Passeriformes. To accomplish this goal, we investigated amount of sleep, sleep time course, and measures of sleep intensity in 6 birds during baseline sleep and during recovery sleep following 6 hours of sleep deprivation.
  
  
  RESULTS
  Continuous (24 hours) EEG and video recordings were used to measure baseline sleep and recovery sleep following short-term sleep deprivation. Sleep stages were scored visually based on 4-sec epochs. EEG power spectra (0.5-25 Hz) were calculated on consecutive 4-sec epochs. Four vigilance states were reliably distinguished based on behavior, visual inspection of the EEG, and spectral EEG analysis: Wakefulness (W), Drowsiness (D), slow wave sleep (SWS) and rapid-eye movement (REM) sleep. During baseline, SWA during D, SWS, and NREM sleep (defined as D and SWS combined) was highest at the beginning of the major sleep period and declined thereafter. Moreover, peak SWA in both SWS and NREM sleep increased significantly immediately following sleep deprivation relative to baseline.
  
  
  CONCLUSIONS
  As in mammals, sleep deprivation in the white-crowned sparrow increases the intensity of sleep as measured by SWA.
authors.affiliation: Neuroscience Training Program, University of Wisconsin-Madison, USA. sgjones2@wisc.edu
authors.forename: Stephany G
authors.initials: SG
authors.lastname: Jones
authors.forename: Vladyslav V
authors.initials: VV
authors.lastname: Vyazovskiy
authors.forename: Chiara
authors.initials: C
authors.lastname: Cirelli
authors.forename: Giulio
authors.initials: G
authors.lastname: Tononi
authors.forename: Ruth M
authors.initials: RM
authors.lastname: Benca
mesh_terms: D000222:Adaptation, Physiological; D000818:Animals; D001143:Arousal / Q000502:physiology; D004569:Electroencephalography; D006706:Homeostasis / Q000502:physiology*; D012815:Signal Processing, Computer-Assisted; D012890:Sleep / Q000502:physiology*; D012892:Sleep Deprivation / Q000503:physiopathology; D012895:Sleep, REM / Q000502:physiology; D046370:Sparrows / Q000502:physiology*; D014741:Video Recording