Journal Abstracts on Circadian Rhythms and Thermoregulation in Reptiles
Compiled by Melissa Kaplan
circadian system of reptiles: a multioscillatory and multiphotoreceptive
Many parameters exhibited by organisms show daily fluctuations that may persist when the organisms are held in constant environmental conditions. Rhythms that persist in constant conditions with a period close to 24 h are called circadian. Although nowadays most research in this field is focused on the molecular and genetic aspects--and therefore mostly on two animal models (Drosophila and mouse)--the study of alternative animal models still represent a useful approach to understanding how the vertebrate circadian system is organized, and how this fascinating time-keeping system has changed throughout the evolution of vertebrates. The present paper summarizes the current knowledge of the circadian organization of Reptiles. The circadian organization of reptiles is multioscillatory in nature. The retinas, the pineal, and the parietal eye (and, possibly, the suprachiasmatic nuclei of the hypothalamus, SCN) contain circadian clocks. Of particular interest is the observation that the role these structures play in the circadian organization varies considerably among species and within the same species in different seasons. Another remarkable feature of this class is the redundancy of circadian photoreceptors: retinas of the lateral eyes, pineal, parietal eye, and the brain all contain photoreceptors.
rhythm of ERG in Iguana iguana: role of the pineal.
In green iguanas, the pineal controls the circadian rhythm of body temperature but not the rhythm of locomotor activity. As part of a program to investigate the characteristics of this multioscillator circadian system, the authors studied the circadian rhythms of the electroretinographic response (ERG) and asked whether the pineal gland is necessary for the expression of this rhythm. ERGs from a total of 24 anesthetized juvenile iguanas were recorded under four different conditions: (a) complete darkness (DD), (b) dim light-dark cycles (dLD), (c) constant dim light (dLL), and (d) pinealectomized in DD. Results demonstrate that the b-wave component of the ERG shows a very clear circadian rhythm in DD and that this rhythm persists in dLL and entrains to dLD cycles. The ERG response is maximally sensitive during the subjective day. Pinealectomy does not abolish the circadian rhythm in ERG, demonstrating that the oscillator responsible for the ERG rhythm is located elsewhere.
of juvenile lizards (Iguana iguana) in a conflict between temperature
regulation and palatable food.
Juvenile green iguanas were placed in a situation of conflict between two motivations: a thermoregulatory drive and the attraction of a palatable bait. To be able to reach the bait (lettuce), they had to leave a warm refuge, provided with standard food, and venture into a cold environment. In experiment 1 the time interval between sessions with bait, ranging from 1 to 8 days, had no effect on the duration of stay on the bait. This result shows that the lettuce was not a necessary food, deprivation of which would have had to be compensated for. In experiment 2 as the ambient temperature at the bait decreased the lizards spent less time feeding on lettuce, and they visited the bait less frequently. This result shows that the lizards traded off the palatability of the bait with the disadvantage of the cold. These findings support the hypothesis that a common currency makes it possible for lizards to compare two sensory modalities.
circadian organization in a vertebrate, Iguana iguana.
The lizard Iguana iguana when kept in constant ambient temperature displays endogenously generated circadian rhythms of body temperature and locomotor activity. Although surgical removal of the parietal eye has only slight effects on overt circadian rhythmicity, subsequent pinealectomy completely abolishes the rhythm of body temperature. However, the rhythm of locomotor activity is only slightly affected by parietalectomy plus pinealectomy. Our results demonstrate that the pineal complex is centrally involved in the generation and control of the circadian rhythm of body temperature but is only marginally involved in locomotor rhythmicity. Plasma melatonin levels are not significantly reduced by parietalectomy, whereas pinealectomy dramatically lowers the level and completely eliminates the circadian rhythm of melatonin in the circulation. Isolated parietal eye, pineal, and retina all synthesize melatonin with robust circadian rhythmicity when maintained for >/=4 d in culture, although in the intact animal all or almost all of the circulating melatonin comes from the pineal. The circadian system of I. iguana is composed of multiple circadian oscillators that reside in different tissues and have specific and different roles.
pineal complex and melatonin affect the expression of the daily rhythm
of behavioral thermoregulation in the green iguana.
Daily variation in the body temperature of the green iguana (Iguana iguana) was studied by telemetry in laboratory photo-thermal enclosures under a 12Light:12Dark (L:D) photoperiod. The lizards showed robust daily rhythms of thermoregulation maintaining their body temperatures (Tb) at higher levels during the day than during the night. Some animals maintained rhythmicity when kept in constant darkness. On light:dark cycles parietalectomy produced only a transient increase of median Tb in the first or second night following the operation. Pinealectomized lizards on the other hand maintained their body temperatures as significantly lower levels during the day and at significantly higher levels during the night than did sham-operated or intact lizards. This effect was apparently permanent, since one month after pinealectomy lizards still displayed the altered pattern. Plasma melatonin levels in intact animals were high during the night and low during the day and were unaffected by parietalectomy. Pinealectomized lizards showed low levels of plasma melatonin during both the day and the night. A daily intraperitoneal injection of melatonin in pinealectomized animals given a few minutes after the light to dark transition decreased the body temperatures selected by the lizards during the night and increased the body temperatures selected during the following day. Control injections of saline solution had no effect. The significance of these results is discussed in relation to the role of the pineal complex and melatonin in the mediation of thermoregulatory behavior.
rhythm of body temperature in an ectotherm (Iguana iguana).
Ectothermic animals regulate their body temperatures primarily by behavioral adjustment in relation to the thermal characteristics of the environment. Several studies have shown that some vertebrate ectotherms may show a daily pattern of body temperature selection when given a choice of environmental temperature. The pattern of body temperature selection free-runs when the animals are kept in constant darkness, demonstrating the existence of circadian regulation. To test whether there might also be a low amplitude circadian rhythm of body temperature itself, we examined the pattern of body temperature and locomotor activity of the lizard Iguana iguana held in a constant environmental temperature. Both variables were recorded for 3 days in a light:dark cycle and then for 10 days in constant dim light (0.1 lux). Under these conditions the body temperature of the lizard oscillates with a circadian period as does the locomotor behavior. These results demonstrate for the first time that ectothermic animals may display physiologically generated circadian rhythms of body temperature similar to those recorded in endotherms. In some animals the circadian rhythms of body temperature and locomotor activity showed different free-running periods, demonstrating that the body temperature rhythm was not caused by locomotor activity and suggesting internal desyncronization of the two rhythms.
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