European hamsters (Cricetus cricetus) show a transient phase of insensitivity to long photoperiods after gonadal regression

Annual rhythms of body weight and reproduction in the European hamster (Cricetus cricetus) are the result of an interaction between seasonal changes in day length (photoperiod) and seasonal changes in the responsiveness of animals to these photoperiods. The present study demonstrates that under natural conditions European hamsters are not able to perceive long photoperiods (i.e., a 16L:8D cycle) before mid-November. This is an important difference to other hamster species, in which regrowth of the gonads can be stimulated by exposure to long photoperiods at any stage of gonadal regression. The experiments also demonstrate the existence of an annual phase of sensitivity to long photoperiods that starts around mid-November and extends until March/April. During this phase of sensitivity, exposure to a long photoperiod (16L:8D) induced gonadal regrowth within 3 wk. Additional experiments with an accelerated photoperiodic lighting regimen indicated that a photoperiod of approximately 13 h is necessary to stimulate gonadal regrowth. Under natural light conditions in Stuttgart (48.46 degrees N), a photoperiod of 13 h is reached by the beginning of April, which fits well with the finding that the majority of animals kept under a natural light:dark cycle had well-developed gonads by the end of April. Nevertheless, these animals showed a rather variable timing of gonadal regrowth, ranging from early January to late April. This is most likely the result of two processes: first, an endogenous mechanism (photorefractoriness) that induces gonadal recrudescence without any photoperiodic information while the animals are still in their hibernation burrows, and second, a direct stimulatory effect of long photoperiods.     

Photoperiodic control of the termination of breeding and the induction ofmoult in House Sparrows Passer domesticus

Two photoperiodic mechanisms controlling gonadal regression in birds have been identified: absolute photorefractoriness, typical of species with short breeding seasons, where gonadal regression occurs spontaneously during long days, and relative photorefractoriness, where a decrease in daylength is required to induce regression. An experiment was designed to test whether these simply represent extremes of one underlying mechanism. Three groups of male House Sparrows Passer domesticus were transferred from a short photoperiod, 8 h of light: 16 h of darkness per day (8L:16D) to long photoperiods of either 18L:6D, 16L:8D or 13L:11D. Gonadal maturation rates were similar in all three groups; gonadal regression and moult began latest in the 13L:11D group. Four additional groups of sparrows were transferred from 8L:16D to 18L:6D and then transferred to either 13L: 11D or 16L:8D prior to, or shortly after, the onset of gonadal regression. The decrease in daylength prior to regression had no effect on the timing of regression but did advance the onset of moult. Decrease in daylength after the onset of regression increased the rate of regression and the rate of moult. Because a decrease in daylength did not affect the timing of regression, the data do not support the hypothesis that absolute and relative photorefractoriness represent extremes of a single underlying photoperiodic control mechanism. The adaptive significance of the effects of decreasing daylength on the rate of regression and moult is discussed.     

Photoperiodic response of the hypothalamo-pituitary-gonad axis in male and female canaries,Serinus canaria

Although the gonadal photoperiodic response and its influence upon the song control system in canaries have been extensively studied, photoperiodic regulation of the GnRH system has not been investigated. To examine the relationship between photoperiod and the reproductive neuroendocrine system in male and female canaries, three groups of canaries were exposed to chronic short days (8L:16D; Phsens), acute long days (18L:6D; Phstim) and chronic long days (also 18L:6D; Phrefr) to induce the reproductive states of photosensitivity, photostimulation, and photorefractoriness, respectively. Brain sections were processed for GnRH immunocytochemistry. The canaries in this study did not demonstrate consistent or uniform responses to different photoperiodic treatments. In males, gonad size varied with photoperiod; Phstim males had larger gonads than either the Phsens or Phrefr males. In contrast, there was no difference between groups in female gonad size as a result of photoperiodic treatment. Brain GnRH cell number, cell size, and fiber density were similar in all groups. The results suggest that canaries are not as obligatory photoperiodic as previously thought (or at least not all varieties of domestic canaries are). This could be a result of many years of domestication, the natural history of the species, phylogenetic constraint, or a combination of these factors.     

Seasonal regulation of reproduction: altered role of melatonin under naturalistic conditions in hamsters

The seasonal reproductive cycle of photoperiodic rodents is conceptualized as a series of discrete melatonin-dependent neuroendocrine transitions. Least understood is the springtime restoration of responsiveness to winter-like melatonin signals (breaking of refractoriness) that enables animals to once again respond appropriately to winter photoperiods the following year. This has been posited to require many weeks of long days based on studies employing static photoperiods instead of the annual pattern of continually changing photoperiods under which these mechanisms evolved. Maintaining Siberian hamsters under simulated natural photoperiods, we demonstrate that winter refractoriness is broken within six weeks after the spring equinox. We then test whether a history of natural photoperiod exposure can eliminate the requirement for long-day melatonin signalling. Hamsters pinealectomized at the spring equinox and challenged 10 weeks later with winter melatonin infusions exhibited gonadal regression, indicating that refractoriness was broken. A photostimulatory effect on body weight is first observed in the last four weeks of winter. Thus, the seasonal transition to the summer photosensitive phenotype is triggered prior to the equinox without exposure to long days and is thereafter melatonin-independent. Distinctions between photoperiodic and circannual seasonal organization erode with the incorporation in the laboratory of ecologically relevant day length conditions.     

Timing of photorefractoriness in the European starling: significance of photoperiod early and late in the reproductive cycle

In European starlings, as in many other birds inhabiting higher latitudes, gonads develop in response to the increasing daylengths in early spring. Later in the year, however, the hypothalamo-pituitary-gonadal axis becomes refractory to the previously stimulatory long photoperiods and the gonads regress in summer. The present study addresses the question of when during the gonadal growth phase photorefractoriness is determined. A 13-h photoperiod induces testicular development and subsequent testicular regression associated with refractoriness in male starlings. An 11-h photoperiod, in contrast, induces only testicular development, and photorefractoriness never develops. When starlings were transferred to an 11-h photoperiod, either 12 or 25 days following exposure to a 13-h photoperiod, their testes developed to full size, but remained large to the end of the experiment, i.e. refractoriness did not develop. The same was even true of most birds in a third group that were transferred to an 11-h photoperiod after 46 days of the 13-h photoperiod, when gonads had developed to near maximal size. These data show that, in contrast to some other species of passerine birds, the onset of photorefractoriness does not become fixed before the testes have undergone considerable development, and that the photoperiodic conditions experienced at the end of the testicular growth phase are still effective in determining the precise time of onset of photorefractoriness. It is suggested that this peculiarity of the starling is related to the fact that its gonadal development begins rather early in spring and, hence, under much shorter photoperiods than the other species studied.     

Photoperiod as a modifying and limiting factor in the expression of avian circannual rhythms

In three species of birds that migrate long distances, the annual rhythms of gonadal activity, molt, and migratory restlessness (Zugunruhe) persist for more than 1 year under certain constant conditions. The most important zeitgeber for these circannual rhythms is the annual cycle of photoperiod, which adjusts the overall period of circannual rhythms to exactly 1 year and also provides for the appropriate adjustment of seasonal activities to the temporal structure of the environment. This is illustrated by results on garden warblers (Sylvia borin) indicating that the longer photoperiods experienced by individuals wintering far south in the African wintering area phase-advance spring migration and the accompanying gonadal development, relative to those of individuals wintering further north. The rate of acceleration is, however, slow enough to prevent a reproductive cycle during the Southern Hemisphere summer. Hence, endogenous circannual components and zeitgeber stimuli constitute a functional entity that provides as a whole for adaptive temporal programming. This idea is further supported by findings in the pied flycatcher (Ficedula hypoleuca), in which a circannual rhythmicity persists only if photoperiod in winter is at least as short as that normally encountered by the species in its wintering grounds slightly north of the equator. In collared flycatchers (Ficedula albicollis), in contrast, rhythmicity continues under much longer photoperiods, consistent with the fact that the wintering area of this species extends to latitudes far south of the equator. It is proposed that the adaptive function of circannual rhythms can be properly understood only if their interactions with environmental factors, particularly those that play a role as zeitgebers, are analyzed in sufficient detail. The biological significance of circannual rhythms may be more apparent in the context of the environmental constraints limiting their expression than in the often rather restricted set of conditions sustaining spontaneous annual cyclicity.     

Control of annual gonadal cycles in Indian songbirds

Abstract

Reproduction is a part of life cycle with great environmental dependence. In contrast to temperate avian species, which mostly breed during summer, the Indian songbirds have more flexible breeding programs and exhibit a spectrum of reproductive strategies with the breeding season scattered all over the year. Control of breeding cycles in the Indian songbirds, therefore, are broadly viewed in light of two strategies (i) birds showing strong photoperiodic component in regulation of reproductive and post-reproductive events (ii) birds that do not exhibit typical photoperiodic regulation indicating the involvement of an inherent rhythm of reproduction. Both circadian and circannual rhythms have been demonstrated to regulate annual gonadal cycles of Indian songbirds. While photoperiod continues to be a predominant proximate factor for timing of breeding in majority of Indian songbirds investigated so far, some studies reveal the role of non photoperiodic cues such as the food availability, temperature, rainfall, etc. in timing/modulating the timing of breeding. The conversion or non-conversion of thyroxine to triiodothyronine may act as a long or short photoperiod signal and may up or downregulate the synthesis and release of GnRH-I in hypothalamus, FSH and LH in anterior pituitary and gonadal steroids in gonads causing gonadal growth or regression, respectively.     

The development of photorefractoriness in termination of the breeding season in the tropical brahminy myna: role of photoperiod

Gonads of brahminy myna (Sturnus pagodarum) spontaneously regress in July/August when the daylength is still stimulatory. Experiments were conducted to investigate if photoperiod was involved in the timing of gonadal regression and if photorefractoriness terminated the breeding season in this species. The observations obtained in the present study clearly show that: i) increasing photoperiods of spring/summer programmed for eventual gonadal regression in the late summer; ii) the birds developed photorefractoriness to all stimulatory daylengths and consequently the breeding season could not be extended by providing more stimulatory photoperiods; and iii) exposure to short daylength treatment failed to overcome the onset of refractoriness in birds after they had attained full gonadal growth and development. These results suggest that refractoriness is a process used by the brahminy myna to terminate the breeding season, and that this species becomes totally photorefractory.     

Interaction between photoperiod and an endogenous seasonal factor in influencing the diel locomotor activity of the benthic polychaete Nereis virens Sars

The locomotor activity of Nereis virens Sars associated with food prospecting was investigated in response to photoperiod and season using an actograph. Experimental animals which had been reared under natural photoperiods were exposed to two constant photoperiodic treatments, LD 16:8 and LD 8:16, in both the autumn and winter and in the absence of tidal entrainment. Autocorrelation analysis of rhythmicity showed that during the autumn, animals under the LD 16:8 photoperiod displayed a strong nocturnal rhythm of activity, whereas animals under the LD 8:16 photoperiod showed only a weak nocturnal activity rhythm. This is believed to represent an autumn feeding cessation that is triggered when the animals pass through a critical photoperiod LD(crit) <12:>12. Later in the winter, however, animals exposed to both photoperiodic treatments showed strong rhythms of foraging activity irrespective of the imposed photoperiod. It is suggested that the autumn cessation may maximize the fitness of N. virens, a spring-breeding semelparous organism, by reducing risk during gamete maturation, while spontaneous resurgence of activity after the winter solstice permits animals that are not physiologically competent to spawn to accrue further metabolic reserves. This response is believed to be initiated by a seasonal (possibly circannual) endogenous oscillator or interval timer.     

Relative photorefractoriness, prolactin, and reproductive regression in a flexibly breeding sonoran desert passerine, the rufous-winged sparrow, Aimophila carpalis

We tested the hypothesis that adult male rufous-winged sparrows, Aimophila carpalis, exhibit relative photorefractoriness. This condition results in partial loss of sensitivity to photoperiod as a reproductive stimulus after prolonged exposure to long photoperiods and is similar to the mammalian condition called photoperiodic memory. Captive birds were exposed either to 8 h of light/16 h of dark per day (8L) or to 16L for 11 weeks and were then exposed either to 8L, 13L, 14L, or 16L. Testicular diameter, plasma luteinizing hormone (LH), and plasma prolactin (PRL) were measured to assess reproductive system activity in response to photostimulation. In free-living birds, testicular diameter, plasma LH, and PRL were compared in birds caught in September in a year when birds were breeding and in a year when birds were not breeding to further evaluate the role of PRL in the termination of seasonal breeding. Testes completely developed after transfer from 8L to 14L or to 16L and partially developed after transfer from 8L to 13L. However, after 11 weeks of 16L exposure, transfer to 14L caused partial regression and transfer to 13L caused complete regression of the testes. Plasma LH increased in all birds that were transferred from 8L to a longer photoperiod. PRL showed a weak response to longer photoperiod treatment and was elevated in birds after chronic 16L exposure in comparison to birds exposed to chronic 8L. These data indicate that male rufous-winged sparrows lose sensitivity to photoperiod after long photoperiod exposure consistent with the relative photorefractoriness and photoperiodic memory models. Lower PRL in birds that developed testes on 13L and 14L compared to birds that regressed testes on 13L and 14L are consistent with the hypothesis that PRL regulates relative photorefractoriness. However, PRL does not appear to regulate interannual differences in the timing of testicular regression.     

Clock gene daily profiles and their phase relationship in the rat suprachiasmatic nucleus are affected by photoperiod

Rhythmicity of the rat suprachiasmatic nucleus (SCN), a site of the circadian pacemaker, is affected by daylength; that is, by the photoperiod. Whereas various markers of rhythmicity have been followed, so far there have been no studies on the effect of the photoperiod on the expression of the clock genes in the rat SCN. To fill the gap and to better understand the photoperiodic modulation of the SCN state, rats were maintained either under a long photoperiod with 16 h of light and 8 h of darkness per day (LD16:8) or under a short LD8:16 photoperiod, and daily profiles of Per1, Cry1, Bmal1 and Clock mRNA in darkness were assessed by in situ hybridization method. The photoperiod affected phase, waveform, and amplitude of the rhythmic gene expression as well as phase relationship between their profiles. Under the long period, the interval of elevated Per1 mRNA lasted for a longer and that of elevated Bmal1 mRNA for a shorter time than under the short photoperiod. Under both photoperiods, the morning and the daytime Per1 and Cry1 mRNA rise as well as the morning Bmal1 mRNA decline were closely linked to the morning light onset. Amplitude of Per1, Cry1, and Bmal1 mRNA rhythms was larger under the short than under the long photoperiod. Also, under the short photoperiod, the daily Clock mRNA profile exhibited a significant rhythm. Altogether, the data indicate that the whole complex molecular clockwork in the rat SCN is photoperiod dependent and hence may differ according to the season of the year.     

Circannual rhythms in birds

In many birds, reproduction, molt, migration and other seasonal activities are controlled by endogenous circannual rhythmicity. Under constant conditions, this rhythm persists for many cycles with a period deviating from 12 months. Whether or not the rhythm is expressed depends on day length (photoperiod), which thus represents an important permissive factor in the process of rhythm generation. In nature, circannual rhythms are usually synchronized by the seasonal changes in photoperiod. However, equatorial birds may use daytime light intensity, which changes with the annual cycle of dry and rainy seasons, as a synchronizing zeitgeber. Photoperiod also modulates the rate of progress of the successive phases of the rhythmicity, such that an optimal adjustment to the annual environmental cycle is guaranteed. Populations of a given species may differ in their responsiveness to photoperiod in a manner that can be described as 'adaptive population-specific reaction norms'. In young migratory songbirds a circannual program determines changes in migratory direction and, at least partly, the time course and distance of migration. This circannual mechanism is replaced or supplemented in older birds by mechanisms formed on the basis of learning and memory. In general, circannual rhythms are intimately involved in the seasonal organization of a bird's behavior, providing the substrate onto which seasonal environmental factors act.     

Circadian rhythmicity and photoperiodism in the pitcher-plant mosquito: adaptive response to the photic environment or correlated response to the seasonal environment?

Many plants and animals use the length of day or photoperiod to cue their seasonal patterns of development, reproduction, dormancy, and migration. Among temperate arthropods, the median or critical photoperiod increases with latitude or altitude. Concomitantly, in beetles, moths, mites, flies, and mosquitoes, there is a declining expression of a rhythmic, presumably circadian-based, component of photoperiodic response. It has been proposed that the long summer days in the north select for a reduced response to light by the circadian clock, which results in this declining rhythmic expression and, consequently, longer northern critical photoperiods. However, these patterns might also be due to direct, seasonal selection on the critical photoperiod itself, which results in a correlated reduction in the rhythmic component as a result of internal physiological constraints within the organism. Using standard light duration and selection experiments, we show that evolution of photoperiodic time measurement in the mosquito, Wyeomyia smithii, results from the direct response of critical photoperiod to seasonal selection and a correlated response of the rhythmic component of photoperiodic time measurement. We conclude that expression of the circadian clock is necessary neither for the central mechanism of photoperiodic time measurement nor for the adaptive modification of critical photoperiod.     

Importance of photoperiodic signal quality to entrainment of the circannual reproductive rhythm of the ewe

An endogenous circannual rhythm drives the seasonal reproductive cycle of a broad spectrum of species. This rhythm is synchronized to the seasons (i.e., entrained) by photoperiod, which acts by regulating the circadian pattern of melatonin secretion from the pineal gland. Prior work has revealed that melatonin patterns secreted in spring/summer entrain the circannual rhythm of reproductive neuroendocrine activity in sheep, whereas secretions in winter do not. The goal of this study was to determine if inability of the winter-melatonin pattern to entrain the rhythm is due to the specific melatonin pattern secreted in winter or to the stage of the circannual rhythm at that time of year. Either a summer- or a winter-melatonin pattern was infused for 70 days into pinealectomized ewes, centered around the summer solstice, when an effective stimulus readily entrains the rhythm. The ewes were ovariectomized and treated with constant-release estradiol implants, and circannual cycles of reproductive neuroendocrine activity were monitored by serum LH concentrations. Only the summer-melatonin pattern entrained the circannual reproductive rhythm. The inability of the winter pattern to do so indicates that the mere presence of a circadian melatonin pattern, in itself, is insufficient for entrainment. Rather, the characteristics of the melatonin pattern, in particular a pattern that mimics the photoperiodic signals of summer, determines entrainment of the circannual rhythm of reproductive neuroendocrine activity in the ewe.     

Photorefractoriness and energy availability interact to permit facultative timing of spring breeding

In seasonally breeding mammals, vernal reproductive development is not directly triggered by increases in day length, rather, an endogenous program of photorefractoriness to short winter days initiates spontaneous development in advance of spring. The transition to the reproductive phenotype is energetically demanding. How food availability in late winter and early spring impacts the onset and expression of photorefractoriness is not known. In this study, male Siberian hamsters were born into a simulated natural photoperiod, and at the winter solstice, they were subjected to a restricted feeding protocol in which a daily food ration was provided in an amount equal to ad libitum (AL) intake during the weeks preceding the solstice. Over the next several months, AL-fed control hamsters exhibited spontaneous recrudescence or spontaneous development. In contrast, vernal reproductive development was abolished in most food-rationed hamsters. In food-rationed hamsters that did exhibit recrudescence, conspicuous delays in the onset of gonadal development and decreases in the magnitude of growth were evident. In all hamsters, the termination of food rationing triggered rapid gonadal development. The data indicate that late winter/early spring increases in environmental food availability are required for the normal manifestation of photorefractoriness-induced reproductive development and suggest that a function of photorefractoriness may be merely to disinhibit the reproductive axis from photoperiodic suppression. Vernal gonadal development or recrudescence appears to be strongly affected by proximate energy availability.     

Is food availability a circannual zeitgeber in tropical birds? A field experiment on stonechats in tropical Africa

Equatorial stonechats (Saxicola torquata axillaris) in Africa are seasonal breeders like their temperate-zone conspecifics (S.t. rubicola). Their annual cycle in gonadal size and function is controlled by an endogenous circannual rhythmicity that has been shown to run for up to 10 years in a constant equatorial photoperiod under laboratory conditions, with a period deviating from 12 months. In nature, however, this rhythm is synchronized with the actual year. Because photoperiod is essentially constant at the equator, it is likely that other environmental factors act as zeitgebers. The authors test whether food availability affects reproductive cycles of free-living East African stonechats. The authors offered supplemental food to the birds 2 months before the regular onset of the breeding season. Supplementally fed males started to sing and display earlier than males of control pairs that did not receive extra food. Although the supplemented food advanced the onset of the breeding season in the pairs that were fed, the onset of the postnuptial molt following the breeding season was not correspondingly shifted. Furthermore, in the year following the experiment, all pairs initiated breeding at the same time. The authors conclude that food availability does not act as a zeitgeber, but rather as a factor that modifies the timing of reproduction without affecting the underlying rhythmicity. The authors propose that this is adaptive under environmental conditions that are relatively constant within a given year but may vary considerably between years. The zeitgeber synchronizing the endogenous rhythmicity remains to be identified.     

Seasonal changes in song nuclei and song behavior in Gambel's white-crowned sparrows

In males of several songbird species, the morphology of forebrain nuclei that control song changes seasonally. The only seasonally breeding songbird in which seasonal changes in the structure of song control nuclei have been reported not to occur is the nonmigratory Nuttall's subspecies of white-crowned sparrow. In the present study, we manipulated photoperiod and plasma testosterone concentrations in captive male white-crowned sparrows of the migratory Gambel's subspeices. Males exposed to photoperiods and plasma testosterone concentrations typical of those experienced by wild breeding males had larger song control nuclei than males held on a winter photoperiod. We also found seasonal change in stereotypy of spectral and temporal parameters of song in wild Gambel's white-crowned sparrows. We hypothesize that seasonal changes in song control nuclei may correlate with seasonal changes in song stereotypy. © 1995 John Wiley & Sons, Inc.     

The contribution of an hourglass timer to the evolution of photoperiodic response in the pitcher-plant mosquito, Wyeomyia smithii

Abstract Photoperiodism, the ability to assess the length of day or night, enables a diverse array of plants, birds, mammals, and arthropods to organize their development and reproduction in concert with the changing seasons in temperate climatic zones. For more than 60 years, the mechanism controlling photoperiodic response has been debated. Photoperiodism may be a simple interval timer, that is, an hourglasslike mechanism that literally measures the length of day or night or, alternatively, may be an overt expression of an underlying circadian oscillator. Herein, we test experimentally whether the rhythmic response in Wyeomyia smithii indicates a causal, necessary relationship between circadian rhythmicity and the evolutionary modification of photoperiodic response over the climatic gradient of North America, or may be explained by a simple interval timer. We show that a day-interval timer is sufficient to predict the photoperiodic response of W. smithii over this broad geographic range and conclude that rhythmic responses observed in classical circadian-based experiments alone cannot be used to infer a causal role for circadian rhythmicity in the evolution of photoperiodic time measurement. More importantly, we argue that the pursuit of circadian rhyth-micity as the central mechanism that measures the duration of night or day has distracted researchers from consideration of the interval-timing processes that may actually be the target of natural selection linking internal photoperiodic time measurement to the external seasonal environment.     

Phenotypic differences in the GnRH neuronal system of deer mice Peromyscus maniculatus under a natural short photoperiod

The neural mechanisms by which short photoperiod induces gonadal regression among seasonally breeding mammals are not well understood. One hypothesis suggests that the proximate cause of seasonal gonadal regression is a photoperiod-induced modification in GnRH secretion. This hypothesis is indirectly supported by our recent findings using immunocytochemistry which identified specific photoperiod-induced adjustments in the number and morphology of GnRH containing neurones between reproductively competent and reproductively regressed laboratory housed male deer mice. Herein, we report that the GnRH neuronal system is similarly affected in reproductively responsive and nonresponsive wild male deer mice Peromyscus maniculatus exposed to a natural short photoperiod. The distribution of immunoreactive (IR)-GnRH neurones was nearly identical in field caught animals and those housed under artificial photoperiod in the laboratory. Compared with reproductively nonresponsive males, reproductively responsive mice from the field population possessed a greater total number of IR-GnRH neurones, a greater number of IR-GnRH neurones within the lateral hypothalamus, and a greater proportion of bipolar IR-GnRH neurones. Each of these distributional and morphological characters was consistent with our findings in laboratory housed male deer mice exposed to an artificial short photoperiod. Taken together, these data underscore the validity of using an artificial photoperiod to evaluate seasonal adjustments in reproductive function in the laboratory.     

The circadian system of ruin lizards: a seasonally changing neuroendocrine loop?

Previous studies have shown that the amplitude of daily melatonin production in cultured ruin lizard pineal organs explanted in the summer is significantly higher than that from organs explanted in the winter. To test whether seasonal photoperiodic changes are decoded autonomously by the pineal gland, pineals explanted in summer were cultured in vitro and exposed to changes between winter and summer photoperiods. The changes in photoperiod duration did not affect the daily profiles of in vitro melatonin production. The discrepancy between the present in vitro results and those from lizards exposed to winter or summer photoperiods before pineal explantation supports the view that circadian information entering the pineal gland via its innervation is involved in determining seasonal changes of melatonin production in ruin lizards. We further examined whether a central component of the circadian system of ruin lizards, specifically the retinae of the lateral eyes, expresses similar seasonal changes in function as does the pineal gland. We did not find any difference between summer and autumn-winter in the effectiveness of either bilateral retinalectomy or optic nerve lesion-at the level of the optic chiasm-in altering circadian locomotor behavior in constant conditions. Both surgical procedures mostly induced a shortening of the free-running period of the locomotor rhythm of similar magnitude in all seasons. Thus, the retinae do not appear to participate in the seasonal reorganization of the circadian system in ruin lizards.