Common features of photoperiodism in plants and animals

Abstract— Light break experiments show the role of the circadian clock in photoperiodic timemeasurement. Additional evidence comes from experiments which show the close relation between temperature- or light-induced phase shifts in cycles of photoperiodic responsiveness and simultaneous phase shifts of other processes which are under control of the circadian clock. Using the circadian clock is a common feature of photoperiodism in plants and animals. This clock controls quantitative and qualitative changes in the responsiveness to light. However, quite different pigment systems may be involved, resulting in different action spectra for light break effects. In many cases, these action spectra differ also from the action spectra for phase shifts in the circadian clock.     

Light Effects on Cyclic Nucleotide Levels and Phase Shifting of the Circadian Clock in Neurospora crassa

Abstract— Mycelia of Neurospora crassa (band [bd] mutant) were exposed to white light (blue light intensity 3.5 μ.mol s^?1 m^?2) of different durations during constant darkness. The concentrations of different second messenger molecules and the phase shifting of the circadian rhythm were determined during light exposures at circadian time (ct) 12 and thereafter. These light exposures elicited 8-12 ct units delay phase shifts but did not change the amount of inositol 1,4,5-triphosphate (InsP₃). In contrast, significant effects of light were observed on cyclic adenosine 3′,5′-monophosphate (cAMP) levels, which increased transiently about 30-90 s after the onset of light. The same kinetics was observed under continued exposure to light as well as after 10 s and 2 s of light followed by darkness. The relative amount of the cAMP-dependent protein kinase A (PKA) in the form of its catalytic sub-units was determined in isolated nuclei of the bd strain in relation to total nuclear proteins by means of western blot analysis, using a heterologous antibody. The nuclear PKA content changed parallel to the cAMP changes. The transient increase of cAMP did not occur in the “blind”white collar mutant (wc-2). Long-term kinetics of cAMP changes after different light pulses in bd showed the initial increase and a return to control levels about 10 min after the onset of the light pulse and a subsequent longer lasting decrease. Light-induced cAMP changes and light-induced phase shifts showed different duration dependencies, thus indicating that cAMP may not play a role in the signal transduction pathway to the clock. Light exposures, furthermore, led to a significant decrease of the cyclic guanosine 3′,5′-monophosphate (cGMP) level. Long-term kinetics of the cGMP content again showed the immediate decrease after 2 min and a slow recovery to (or above) control values after several hours. Various calcium channel blockers (nickel, cobalt, nifedipine, dantrolene, lanthanum) and the calcium calmodulin inhibitor chlorpromazine did not affect the phase shifting by light nor did they significantly phase shift the circadian rhythm in the dark themselves (again determined at ct 12). The data showed that InsP₃ did not change after exposure to light, whereas cAMP increased and cGMP decreased. There was no evidence, however, that these changes play a role in light signal transmission to the clock.     

Light effects in yeast: persisting oscillations in cell division activity and amino acid transport in cultures of Saccharomyces cerevisiae entrained by light-dark cycles.

Abstract— In addition to the direct inhibitory effects of visible light (cool-white fluorescent, < 3500 lux) on growth and amino acid transport previously reported for cultures of the yeast Saccharomyces cereui-siae (strain Y185 rho⁺) grown at 12°C in medium containing glucose, yeast carbon base, KH₂PO₄, ammonium ion and proline, we have found evidence for endogenous, light-entrainable, self-sustaining circadian and ultradian oscillators underlying both cell division and transport activity. Diurnal light (? 3000 lux) cycles (LD), imposed on cultures previously grown in the dark, phased or synchronized cell number increase to a 24-h period with bud release being confined primarily to the dark intervals (although not necessarily every cell divided during any given division ‘burst’). The observed division or budding rhythm freeran with a circadian period (?26h) only approximating 24 h for a number of days in constant darkness (DD) following prior entrainment by LD, thereby implicating an endogenous circadian clock. Further, a similar light-entrainable circadian rhythm in the uptake of “C-histidine or ¹4C-lysine occurred in nondividing (or very slowly dividing) cultures during the “stationary” (infradian) phase of growth synchronized by a 24-h LD cycle and then released into DD for as long as 10 days. Some experiments revealed a bimodal (ultradian) periodicity in both LD and DD with secondary peaks or shoulders occurring at intervals of ?12h, corresponding approximately to subjective ‘dawn’ and ‘dusk’. Transport in cultures of the Y185 rho^- petite mutant, which lacks cytochromes a/a₃, b and c_l, could not be synchronized by LD cycles, a finding that is consistent with the hypothesis that these chromophores may be primary photoreceptors for synchronization of rhythms in this microorganism.     

EXTRARETINAL PHOTORECEPTION IN LIZARDS

Abstract— In lizards both photoperiodic photoreception and entrainment (synchronization) of the biological clock by environmental light cycles can be accomplished by extraretinal photoreceptors. Testicular size and spermatogenic activity in the green anole, Anolis carolinensis , is maintained under long (14 h) photoperiods in the summer in both sighted and blinded lizards whereas short days (6h) cause testicular regression. In the fall, long photoperiods cause testicular recrudescence and maturation in blinded and blinded-parietalectomized anoles as well as in sighted anoles.
Extraretinal photoreceptors can mediate entrainment of the circadian locomotor rhythm in every lizard species examined to date (8 species). The photosensory pineal complex in lizards (pineal organ and parietal eye) may not be involved in entrainment; in any case, its removal does not prevent entrainment of blinded lizards. Localization experiments have shown that the brain is the site of the extraretinal receptors mediating entrainment. The lateral eyes can also contribute to entrainment. In some species, but not all, blinding alters the pattern of the entrained activity rhythm. The free-running activity rhythms of two species of lizards exposed to continuous illumination respond very differently to blinding; these experiments yield insight into the mechanisms of entrainment in the two species as well as showing that caution must be observed when generalizing about the role of retinal and extraretinal receptors not only among different vertebrate classes but also among species of the same class.     

MULTIPLE ACTION OF FAR-RED LIGHT IN PHOTOPERIODIC INDUCTION and CIRCADIAN RHYTHMICITY

Abstract— It is generally accepted that phytochrome influences the photoperiodic induction of flowering through its interaction with the circadian clock mechanism. We have attempted to separate the effects of phytochrome on the clock mechanism from those that mediate flowering directly by examining a number of responses that are unrelated to flowering but are also regulated by the circadian clock. Gas exchange measurements of both CO₂ and H₂0 vapor were monitored under light conditions (200 μmol m ² s⁻¹) where the addition of far-red energy is required for the maximal promotion of flowering. In addition, photosynthetic capacity and maximal transpiration rates were measured in plants grown under continuous dim (20 μmol m⁻² S⁻') light, with or without supplemental far-red, by exposing them briefly to saturating fluxes (1000 μmol m⁻² s^-l) of light. Net CO₂ fixation was very weakly rhythmic in plants grown under both high and low light and this weak oscillation was completely suppressed by far-red light. Far-red also suppressed the rhythm in transpiration under high light, but the rhythm was immediately reinstated when the far-red light was removed. The phase of this rhythm was also reset with the next peak always occurring15–18 h after the far-red was turned off. When grown under dim light, the transpiration rhythm was not suppressed and the amplitude of the oscillation was more than doubled. Far-red light appears to interact with the rhythm in transpiration in a manner suggesting that the stomatal rhythm may be coupled to the same clock oscillator that regulates the flowering rhythm.     

CIRCADIAN RHYTHM OF Robinia pseudoacacia LEAFLET MOVEMENTS: ROLE OF CALCIUM AND PHYTOCHROME

Abstract— The effect of external calcium level, calcium ionophore A23187 and red light on the circadian rhythm of Robinia pseudoacacia leaflet movements has been studied. Fifteen minute red light pulses shifted the phase of leaflet rhythmic movement with a phase-response curve type 0. Maximum advances and delays (about 10 h and 8 h, respectively) were obtained between circadian time (CT) 10 and CT 12 at the end of a subjective day. An almost null effect was obtained at the end of a subjective night. Phytochrome is the photoreceptor involved in phase shifting since this effect of red light is reversed by 5 min of far red light. Two hour pulses of external calcium, applied as CaCl₂ (10 m M ), and 2 h pulses of calcium ionophore A23187 (10–50 μM) also shifted the phase of leaflet circadian movement and caused the same type of phase-response curve, with maximum advances and delays at the same time as those produced by red light. Two hour pulses of an external calcium chelator, EGTA (5 m M ), and a calcium channel blocker, LaCl₃ (10–50 m M ), damped the circadian rhythm or did not change the phase when they were applied at lower concentration. These results indicate that phytochrome could control the circadian oscillator, which drives Robinia leaflet movements by increasing the intracellular calcium concentration.     

BLUE LIGHT PHOTORECEPTION IN Neurospora ORCADIAN RHYTHM: EVIDENCE FOR INVOLVEMENT OF THE FLAVIN TRIPLET STATE

The mechanism of the photoreceptor acting on the circadian conidiation rhythm of Neurospora crassa was studied, with the following results: (1) the efficiency of 8-haloflavins as sensitizers increased with their triplet yields. (2) Phase shifts were not abolished by removal of oxygen prior to illumination. (3) Oxygen inhibited phase shifts when introduced into the cultures after light treatment. It is proposed that the blue light photoreceptor for the circadian clock of Neurospora crassa acts (1) from its triplet state, but (2) not via singlet oxygen; (3) signal transduction involves (an) oxygen-sensitive intermediate(s).     

Nonvisual Opsins and the Regulation of Peripheral Clocks by Light and Hormones

The molecular clock machinery is conserved throughout evolution. However, how environmental cues are perceived has evolved in such a way that peripheral clocks in mammals require a variety of signals, including hormones. On the other hand, in nonmammalian cells able to directly detect light, light seems to play a major role in the synchronization of the clock. The interaction between perception of circadian light by nonvisual opsins and hormones will be discussed under the perspective of clock synchronization at the molecular level.     

EFFECTS OF PHASIC AND TONIC LIGHT INPUTS ON THE ORCADIAN ORGANIZATION OF THE SQUIRREL MONKEY

Abstract— The organization of the circadian timing system in Saimiri sciureus was probed using the phasic (abrupt transition) and tonic (continuous action) effects of light intensity. The behavior of the simultaneously monitored circadian rhythms of feeding behavior, colonic temperature, and urinary potassium excretion was studied in response to the phasic effects of (a) an abrupt 8-h phase delay in the light–dark (LD) cycle and (b) a series of non-24 h LD cycles ( T = 18 to 30 h). These studies demonstrated that the feeding and temperature rhythms were more tightly coupled to the light-dark cycle than was the rhythm of urinary potassium excretion. The tonic effects of constant levels of illumination confirmed this conclusion. In constant light, internal desynchronization spontaneously occurred in 25% of animals with the potassium rhythm exhibiting a period quite different from that of the feeding and colonic temperature rhythms. Thus, the response of the internal circadian timekeeping system to phasic and tonic light inputs shows that the system in this species comprises at least two potentially independent oscillators with differential light sensitivities.     

Crayfish Procambarus clarkii retina and nervous system exhibit antioxidant circadian rhythms coupled with metabolic and luminous daily cycles

Based on previous work in which we proposed midgut as a putative peripheral oscillator responsible for circadian reduced glutathione (GSH) crayfish status, herein we investigated the retina and optic lobe-brain (OL-B) circadian GSH system and its ability to deal with reactive oxygen species (ROS) produced as a consequence of metabolic rhythms and light variations. We characterized daily and antioxidant circadian variations of the different parameters of the glutathione system, including GSH, oxidized glutathione (GSSG), glutathione reductase (GR) and glutathione peroxidase (GPx), as well as metabolic and lipoperoxidative circadian oscillations in retina and OL-B, determining internal and external GSH-system synchrony. The results demonstrate statistically significant bi- and unimodal daily and circadian rhythms in all GSH-cycle parameters, substrates and enzymes in OL-B and retina, as well as an apparent direct effect of light on these rhythms, especially in the retina. The luminous condition appears to stimulate the GSH system to antagonize ROS and lipid peroxidation (LPO) daily and circadian rhythms occurring in both structures, oscillating with higher LPO under dark conditions. We suggest that the difference in the effect of light on GSH rhythmic mechanisms of both structures for antagonizing ROS could be due to differences in glutathione-system coupling strength with the circadian clock.     

EXTRARETINAL PHOTORECEFTION IN INSECTS

Abstract— Extraretinal photoreceptors are widespread among insects and function in the photoperiodic control of development and in the entrainment of circadian rhythms. The effects of light on the daily and seasonal regulation of brain neuroendocrine activity are mediated solely through extraretinal photoreceptors. In primitive insects, the eyes participate in the entrainment of nonendocrine circadian rhythms such as the locomotor rhythm. In more advanced forms, however, extraretinal pathways appear to be the only pathway for the entrainment of all rhythms thus far examined. But even in this latter case, the eyes sometimes effect a masking of the expression of the overt rhythm. An exact localization of the extraretinal receptors has not been accomplished, but in all studies to date they appear to be associated with the cerebral lobe region of the brain. Action spectra for photoperiodic responses have been determined for a number of insects. In general the responses are maximally sensitive in the blue with a marked decline in the red although exceptions do exist. Complete action spectra for circadian responses have been determined only for two insects. In both cases a plateau of sensitivity extends through the blue with a steep drop at longer wavelengths. From the action spectra data, the extraretinal receptors appear to have a threshold sensitivity less than 3 times 10^-2 J/m². The pigment nature of the receptor is unknown although it appears not to be a carotenoid derivative.     

EXTRARETINAL PHOTORECEPTION IN ENTRAINMENT OF CRUSTACEAN ORCADIAN RHYTHMS*

Abstract— An attempt was made to determine whether entrainment of the circadian rhythms of locomotor activity and ERG amplitude of the crayfish involved extraretinal photoreception. The results of a variety of experiments involving surgical lesions and localized illumination provided evidence that both rhythms can be entrained via an extraretinal pathway. The data also demonstrate that the caudal photoreceptor is unnecessary for entrainment. Our evidence to date suggests that the extraretinal photoreceptor is located in the supraesophageal ganglion; however, the possibility of photoreceptive input from other regions of the CNS, particularly the optic lobe, has not been eliminated. It is also tentatively concluded that the circadian oscillators for both the locomotor and the ERG amplitude rhythms are located within the supraesophageal ganglion, but more data are needed to confirm this conclusion.     

Regulation of the Cyanobacterial Circadian Clock by Electrochemically Controlled Extracellular Electron Transfer

There is growing awareness that circadian clocks are closely related to the intracellular redox state across a range of species. As the redox state is determined by the exchange of the redox species, electrochemically controlled extracellular electron transfer (EC‐EET), a process in which intracellular electrons are exchanged with extracellular electrodes, is a promising approach for the external regulation of circadian clocks. Herein, we discuss whether the circadian clock can be regulated by EC‐EET using the cyanobacterium Synechococcus elongatus PCC7942 as a model system. In vivo monitoring of chlorophyll fluorescence revealed that the redox state of the plastoquionone pool could be controlled with EC‐EET by simply changing the electrode potential. As a result, the endogenous circadian clock of S. elongatus cells was successfully entrained through periodically modulated EC‐EET by emulating the natural light/dark cycle, even under constant illumination conditions. This is the first example of regulating the biological clock by electrochemistry.     

Daily and circadian variation in survival from ultraviolet radiation in Chlamydomonas reinhardtii

The survival of organisms depends on their ability to adapt to their environment, one important aspect of which is the daily cycle of day and night. During the day, organisms use a variety of strategies to protect themselves from deleterious ultraviolet (UV) wavelengths of sunlight. Among those strategies could be timing of UV-sensitive cellular processes to occur at night to avoid UV-induced damage. We tested whether the unicellular alga Chlamydomonas reinhardtii uses this strategy by measuring the survival of cells following exposure to UV radiation at different phases of the day. Chlamydomonas cells displayed a rhythm of survival from UV radiation where the most sensitive phases occurred during the end of the day and at the beginning of the night. This phase of sensitivity corresponds to the time of nuclear division. The rhythm continues in constant light indicating control by a circadian clock. The results presented here suggest a hypothesis of how circadian clocks may have evolved; a temporal program whereby light-sensitive processes are timed to avoid sunlight-induced damage would be advantageous and therefore selected.     

ENTRAINMENT OF HUMAN ORCADIAN RHYTHMS BY LIGHT-DARK CYCLES: A REASSESSMENT

Abstract— Man is the only eukaryotic organism in which it has been reported that the circadian system cannot be entrained to a 24-h period by a simple light-dark (LD) cycle. In this paper, we reexamine the evidence for that claim and demonstrate that there were some fundamental flaws in the experimental design of the previous studies on which this conclusion was based. We report new studies in which we tested the efficacy of LD cycles in entraining the circadian rhythms of human subjects living in isolation from environmental time cues. We found that the cyclic alternation of light and dark, when applied to human subjects in a comparable way to experiments in other species, was an effective entraining agent. Our results and a critical review of the literature indicate that a LD cycle alone can be an effective environmental synchronizer of the human circadian timing system. Other factors, such as the knowledge of time of day, social contacts, the feeding schedule, and the imposed rest-activity schedule may contribute to stable entrainment, although their relative strengths as synchronizers have yet to be determined.     

CIRCADIAN PHASE OF SPARROWS: CONTROL BY LIGHT AND DARK

Abstract— Sparrows ( Passer domesticus ) are day-active birds which exhibit circadian rhythms of perch-hopping activity. The phases of sparrow's circadian rhythms were studied following single 4 h light pulses, single 4 h dark pulses, doublet treatments of light and dark pulses, and a 10 h light pulse.
The sparrows exhibited a phase response curve to 4 h light pulses with maximum phase advances (3.8 h) at CT20 and maximum phase delays(–1.3 h) at CT16. The sparrows also displayed a phase response curve to dark pulses with maximum phase advances (2.2 h) at CT16 and maximum phase delays at CTO(–0.7 h).
The remaining pulses were imposed during the subjective dark-time. The 10 h pulse beginning 1 h after lights-out produced a 2.2 h phase shift. The doublet of 2 h pulses that were the "skeleton" of the 10 h pulse produced a 2.5 h phase shift. The early 2 h pulse, applied by itself resulted in a -0.4 h delay; the late 2 h pulse applied singly produced a 3.1 h advance. When an early 3 h dark pulse was imposed together with a late light pulse, the phase was advanced 3.6 h; singly the pulses produced 1.8 h and 2.7 h advances.     

On the Role of Exponential Smoothing in Circadian Dosimetry

The effects lighting has on health through modulation of circadian rhythms are becoming increasingly well documented. Data are still needed to show how light exposures are influenced by architecture and lighting design and circadian dosimetry analyses should provide duration, phase and amplitude measures of 24 h exposure profiles. Exponential smoothing is used to derive suitable metrics from 24 h light measurements collected from private dwellings. A further application of these modified exposure time series as physiological models of the light drive is discussed. Unlike previous light drive models, the dose rate persists into periods of darkness following exposures. Comparisons to long duration exposure studies suggest this type of persistent light drive model could be incorporated into contemporary physiological models of the human circadian oscillator.     

EFFECTS OF INORGANIC NITROGEN ON THE RESPONSE OF LEMNA CARBON DIOXIDE OUTPUT TO LIGHT QUALITY AND TIMING*

Abstract —The effects of various light/dark schedules on the time course of CO₂ output by axenic cultures of the short-day plant Lemna perpusilla 6746 differ substantially depending on whether the medium is N-less or contains NH₄ or NO₃ as the sole N source. The steady-state pattern achieved with a daily 1/4 h light pulse in N-less medium is essentially the same whether the light is red or far-red; on NO₃ or NH₄, however, the red and far-red patterns differ in form and suggest the action of a ‘Pfr-hourglass’ timer. In darkness, following either continuous light or entrainment to kh red light daily, CO₂ output oscillates for three or more circadian cycles on NH₄ medium and for at least two on N-less, but damps after a single cycle on NO₃. A schedule of 1/4 h red light every 12 h elicits a 24 h periodicity on NO₃ or NH₄ media and a 12 h periodicity on N-less medium, while a similar far-red schedule elicits a 12 h periodicity on all three. CO₂ output patterns on each of the media respond differently to varying the daily span of light from 1/4 to 6 to 12 h. These results are probably due to differential effects of changing N status on the proportion of total C O₂ arising from various metabolic reactions. They suggest that, rather than being a simple, unitary indicator, CO₂ output can be made to reflect different processes on different media, increasing its value as a real-time indicator of events underlying photoperiodism.