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Spontaneous Blink Rates of Birds 92 SHORT COMMUNICATIONS- Condor 85:92-93 in daylight, but in fluorescentdim blue light in the case 8 The Cooper Ornithological Souety 1983 ofthe ScreechOwl. Observationswere made either at close range or with a spotting scope. After the bird had habit- SPONTANEOUS BLINK RATES uated to us (1-5. min), we counted the number of times OF BIRDS that it blinked in 10 separate I-min periods while it sat quietly. For most species,we used at least two individuals to determine a mean blink rate. Both bilateral and uni- lateral blinks aswell asclosure ofthe nictitating membrane SUZANNE J. KIRSTEN were counted, but are not distinguishedin this report. AND The blink rates of most individuals and specieswere relatively constant under quiet, unchanging test condi- EDWARD B. KIRSTEN tions. Mean values ranged from 1.5/min for the Screech Owl to more than 50/min for the Rock Dove and Ruby- throated Hummingbird. In several species,including the The blinking activity of birds has received little attention White-winged Dove, Wild Turkey, and Rock Dove, blink- in the scientific literature, although the subject has been ing appeared to be correlated with head movement, sug- studied in more depth in mammals (Wood and Saunders gestingthat the vestibular apparatusmodulates the blink 1962, Zametkin et al. 1979). Mammals have characteristic reflex in thesebirds, as has been reported in humans (Pon- species-specificblink rates that are independent of light, der and Kennedy 1928). heat, or humidity (Blount 1927, Ponder and Kennedy Although we did not control the amount of light to 1928). They range in frequency from 0-4/min in the rat, which the birds were exposed,the blink ratesdiffered little mouse, hamster, and fox, to 8-22/min in primates, to 30- between individuals in bright light and those in the shade. 36/min in the domestic pig (Hall 1945, Stevens and Liv- Indeed, a blind ScreechOwl blinked at a mean rate (1.2 ermore 1978). Moreover, the blink rate of diurnally active blinks/min) that did not differ significantly from that of mammals is more than 10 times higher than that of noc- conspeciticswith normal vision (1.5 blinks/min). This ob- turnally active species. In this study, we measured the servation suggeststhat in birds, as in humans, the blink spontaneousblink rates of 25 species of birds and ex- rate for individual speciesdoes not depend upon photo- amined them for similar diurnal/nocturnal patterns. Ac- stimulation of the retina, provided the light to which sub- curate data on blink rate could provide a way of assessing jects are exposed is not extremely bright. the physiologicalcondition of individual birds. Altema- Our resultsalso suggestthat diurnally active birds blink tively, if suchrates increasewith ageas they do in humans more frequently than nocturnally active species(Table 1). (Zametkin et al. 1979) blinking may provide an estimate In general, all speciesof owls blinked less than 10 times of age, much as external morphological features are used per minute, whereasdiurnal specieshad higher mean rates. in certain species. The Snowy Owl, which is active in daytime, had the high- We determined the spontaneousblink ratesof 25 species est rate among the owls. of birds in zoos and bird sanctuaries(Table 1). Our mea- Diurnal mammals also have considerablyhigher blink surementswere made between 09:OOand 15:00 generally rates than nocturnal mammals; furthermore, their rapid TABLE 1. Spontaneousblink rate of birds. Average* Range of n blink rate bbnk rates Nocturnal species Black-crownedNight-Heron (Nycticorax nycticorax) 2 3.2 f 0.8 2-4 Eastern Screech-Owl (Otus asio) 2 1.5 f 0.7 o-3 Great Homed Owl (Bubo virgin&us) 7 3.7 & 2.5 o-7 Snowy Owl (Nyctea scandiaca) 3 9.0 i 2.7 6-12 Burrowing Owl (Athene cunicularia) 2 2.9 i 1.2 o-5 Barred Owl (Strix vuria) 4 5.9 i 3.5 2-11 Northern Saw-whet Owl (Aegohus acadicus) 3 4.0 i 1.1 2-5 Diurnal species Scarlet Ibis (Eudocimus ruber) 3 41.3 f 5.3 33-50 Roseate Spoonbill (Ajaia ajaja) 5 32.5 +- 4.5 24-44 Mallard (Anus plutyrhynchos) 2 28.3 f 3.1 22-32 Barrow’s Goldeneye (Bucephala islandica) 2 40.1 f 6.7 32-5 1 Northern Goshawk (Accipiter gentihs) 1 12.7 & 1.6 10-15 Red-shouldered Hawk (Buteo line&us) 1 11.8 i 1.6 9-14 Swainson’s Hawk (Buteo swainsonz) 3 16.8 i 2.4 12-19 Harris’ Hawk (Purubuteo unicinctus) 1 13.2 * 1.6 lo-16 Golden Eagle (Aquilu chrysaetos) 4 16.8 * 3.5 1 l-22 Bald Eagle (Haliaeetus leucocephalus) 1 21.6 i 5.1 13-27 Crested Caracara (Polyborus plancus) 1 13.5 f 2.6 11-19 Wild Turkey (Meleagris gallopavo) 3 43.3 f 7.9 31-51 Common Snipe (GaNinago gullinago) 1 46.9 ?I 4.4 40-54 Laughing Gull (Larus atricillu) 4 41.0 * 3.7 36-48 Black Skimmer (Rynchops niger) 1 18.8 i 2.9 14-23 Rock Dove (Columba livia) 5 52.0 i 22.1 23-88 White-winged Dove (Zenaidu asiatica) 4 28.0 F 5.8 19-36 Ruby-throated Hummingbird (Archilochus colubris) 1 53.4 i 4.6 48-60 * Values m the table are average rates per minute i SD. The latter refers to the variancr among measurements on mdiwdual birds when n is l-2. SHORT COMMUNICATIONS 93 eye movement (REM) sleep each night is shorter (Zepehn have been shownin this study,their functional significance and Rechtschaffen 1974, Zametkin et al. 1979). We at- must await further investigation. tempted to relate REM sleep to spontaneousblinking in We thank Leonard Saucy of the New Jersey Raptor the specieswe studied, but with little successbecause so Association and Charles Leek of Rutgers University for little is known about the REM sleepof birds. Nonetheless, their assistance. three reports suggestthat owls (Berger and Walker 1972) LITERATURE CITED have episodesof REM sleep that are shorter (5% of total sleeutime) than those of Rock Doves (Walker and Berger BENOIT,J. 1964. The role of the eye and of the hypo- 1972) and’ two falconiforms (Rojas-Ramirez and Tauber thalamus in the photostimulation of gonads in the 1970).,/ all of which are diurnal and have neriods of REM duck. Ann. N.Y. Acad. Sci. 117:204-215. sleep averaging 7-10% of their total sleep time. This re- BERGER,R. J., AND J. M. WALKER. 1972. Sleep in the lationship between diurnal and nocturnal forms is oppo- Burrowing Owl (Speotytocunicularia hypugaea).Be- site to that reported for mammals. The limited data avail- hav. Biol. 7: 183-l 94. able and the primitive natureof REM sleepin birds (Berger BLOUNT,W. P. 1927. Studies of the movements of the and Walker 1972) may, however, preclude a significant eyelids of animals: blinking. Q. J. Exp. Physiol. 8: correlation. 111-125. Phasic regulation of neuroendocrine,metabolic, repro- CRAWFORD, M. L. J., AND R. E. MARC. 1976. Light trans- ductive and motor activity by light is well known in birds. mission of cat and monkey eyelids. Vision Res. 16: For example, in the Mallard exposureof retinal receptors 323-324. responsiveonly to red or orange light stimulated gonadal HALL, A. 1945. The origin and purposesof blinking. Br. growth (Benoit 1964). Similarly, under natural conditions, J. Ophthalmol. 29:445-467. the foraging and reproductive activities of many avian PONDER, E., AND W. P. KENNEDY. 1928. On the act of speciesare closely related to daily and seasonalchanges blinking. Q. J. Exp. Physiol. 18:89-l 10. in photoperiod. While theselong-term effectsof light have ROJAS-RAMIREZ, J. A., AND E. S. TAUBER. 1970. Para- been defined, the function of the brief pulsesof light pro- doxical sleep in two speciesof avian predator (Fal- duced by spontaneousblinking has not. coniformes). Science 167:1754-1755. The primary function of spontaneousblinking is gen- STEVENS, J. R., AND A. LIVERMORE. 1978. Eye blinking erally consideredto be the cleansingand irrigation of the and rapid eye movement: pulsed photic stimulation eyes. Ponder and Kennedy (1928) however, noting that of the brain. Exp. Neurol. 60:541-556. its rate was remarkably constantand independent of such WALKER, J. M., AND R. J. BERGER. 1972. Sleep in the factors as light, temperature, humidity, and the integrity domestic pigeon(Columba livin). Behav. Biol. 7:195- of cranial nerves II through VI, concluded that blinking 203. is centrally mediated and that it servesfunctions beyond WOOD,L. A., AND J. C. SAUNDERS. 1962. Blinking fre- simple cleansing. Blinking causesregular brief exposure quency: a neurophysiologicalmeasurement of psy- of the retina to external light through the red/orange filter chologicalstress. Dis. Nerv. Syst. 23: 158-l 62. of the closed lids or nictitating membrane (Crawford and ZAMETKIN, A. J., J. R. STEVENS, AND R. PITTMAN. 1979. Marc 1976). Eyeblinkshave been shownto evoke electrical Ontogenyof spontaneousblinking and of habituation potentialsand multiple neuronal activity in the visual cor- of the blink reflex. Ann. Neurol. 5:453-457. tex and subcorticalnuclei of cats (Stevensand Livermore ZEPELIN, H., AND A. RECHTSCHAFFEN. 1974. Mammalian 1978). Such periodic interruption of the light striking the sleep, longevity, and energy metabolism. Brain Be- retina may provide the avian central nervous systemwith hav. Evol. 10:425-475. a constant source of pulsed stimulation with which to couplesuch cyclic eventsas the releaseofbiogenic amines, 145 BeechStreet, Nutley, New Jersey071 IO. Received 18 endocrine function, or motor activity. While differences December 1981. Final acceptance3 August 1982. in blinking rate between nocturnal and diurnal species Condor 85:93-95 0 The CooperOrnithological Society 1983 NEST PREDATION IN INSULAR extirpation lists) and Willis (1974) suggestedthat preda- AND MAINLAND LOWLAND tion may be a factor in the loss of some of these species.
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