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PubMed2005

Photic input pathways that mediate the Drosophila larval response to light and circadian rhythmicity are developmentally related but functionally distinct.

The Drosophila melanogaster larval photosensory organ that mediates the response to light consists of bilaterally symmetrical clusters of 12 photoreceptors. These are distinguished on the basis of expression of the rhodopsins Rh5 and Rh6. The Rh6-expressing cells correspond to the Hofbauer-Buchner (H-B) eyelet found later in the posterior margin of the adult compound eye and recently shown to function as an input pathway in the entrainment of circadian rhythmicity in adult Drosophila. In addition, the axons of the larval photoreceptors are found in intimate association with a subset of the main circadian pacemaker neurons located in the developing accessory medulla, the small ventral lateral neurons (LNv). The observed spatial overlap between components of the circadian circuitry, input pathway, and pacemaker neurons-and the larval visual organ-suggest a functional relationship between these two photosensory input pathways. In this study we determined the requirement of specific rhodopsin-expressing photoreceptors including the presumptive H-B eyelet and pacemaker neurons in the larval locomotory response to visual stimuli. Our results demonstrate that two of the most important components of the neuronal circuitry underlying circadian rhythmicity in Drosophila, namely, the extraretinal H-B cluster and the circadian pacemakers, while in intimate association with the larval visual system are not required for the larval motor response to light.

Animals↗
PubMed2006

Central projections of melanopsin-expressing retinal ganglion cells in the mouse.

A rare type of ganglion cell in mammalian retina is directly photosensitive. These novel retinal photoreceptors express the photopigment melanopsin. They send axons directly to the suprachiasmatic nucleus (SCN), intergeniculate leaflet (IGL), and olivary pretectal nucleus (OPN), thereby contributing to photic synchronization of circadian rhythms and the pupillary light reflex. Here, we sought to characterize more fully the projections of these cells to the brain. By targeting tau-lacZ to the melanopsin gene locus in mice, ganglion cells that would normally express melanopsin were induced to express, instead, the marker enzyme beta-galactosidase. Their axons were visualized by X-gal histochemistry or anti-beta-galactosidase immunofluorescence. Established targets were confirmed, including the SCN, IGL, OPN, ventral division of the lateral geniculate nucleus (LGv), and preoptic area, but the overall projections were more widespread than previously recognized. Targets included the lateral nucleus, peri-supraoptic nucleus, and subparaventricular zone of the hypothalamus, medial amygdala, margin of the lateral habenula, posterior limitans nucleus, superior colliculus, and periaqueductal gray. There were also weak projections to the margins of the dorsal lateral geniculate nucleus. Co-staining with the cholera toxin B subunit to label all retinal afferents showed that melanopsin ganglion cells provide most of the retinal input to the SCN, IGL, and lateral habenula and much of that to the OPN, but that other ganglion cells do contribute at least some retinal input to these targets. Staining patterns after monocular enucleation revealed that the projections of these cells are overwhelmingly crossed except for the projection to the SCN, which is bilaterally symmetrical.

Animals↗
PubMed1995

Nitric oxide synthase in tiger salamander retina.

Previous studies have indicated that nitric oxide, a labile freely diffusible biological messenger synthesized by nitric oxide synthase, may modulate light transduction and signal transmission in the retina. In the present work, the large size of retinal cells in tiger salamander (Ambystoma tigrinum) allowed the utilization of nicotinamide adenine dinucleotide phosphate (NADPH)-diaphorase histochemistry and nitric oxide synthase immunocytochemistry to delineate the cell-specific intracellular localization of nitric oxide synthase. NADPH-diaphorase activity was highly concentrated in the outer retina, in rod and cone inner segment ellipsoids, and between and adjacent to the photoreceptor cell bodies in the outer nuclear layer. Examination of enzymatically isolated retinal cells indicated that outer nuclear layer NADPH-diaphorase activity was localized to the distal processes of the retinal glial (Müller) cells and to putative bipolar cell Landolt clubs. Less intense NADPH-diaphorase activity was seen in the photoreceptor inner segment myoid region, in a small number of inner nuclear layer cells, in cap-like configurations at the distal poles of cells in the ganglion cell layer and surrounding ganglion cell layer somata, and in punctate form within both plexiform layers, the pigment epithelium, and the optic nerve. Nitric oxide synthase-like immunoreactivity was similarly localized, but was also concentrated along a thin sublamina centered within the inner plexiform layer. The potential for nitric oxide generation at multiple retinal sites suggests that this molecule may play a number of roles in the processing of visual information in the retina.

Animals↗
PubMed2002

Circadian efferent input to Limulus eyes: anatomy, circuitry, and impact.

Much is known about the anatomy of Limulus retinal efferent neurons and the structural and functional consequences of their activation. Retinal efferent axons arise from cell bodies located in the cheliceral ganglia of the brain, and they project out all of the optic nerves. Their unique neurosecretory-like terminals contact all cell types in lateral eye ommatidia, the retinular cells of the median eye, and the internal rhabdom of ventral photoreceptors. Lateral and median rudimentary photoreceptors are also innervated. The activity of the efferents is circadian. They are active during the subjective night and inactive during the subjective day. Activation of the efferents drives dramatic and diverse changes in the structure and function of Limulus eyes and causes the sensitivity and responsiveness of the eyes to light to increase at night. Relatively little is known about the molecular mechanisms that produce these structural and functional changes, but one efferent-activated biochemical cascade has been identified. The biogenic amine octopamine is released from efferent terminals, and an octopamine-stimulated rise in cAMP in photoreceptors, with a subsequent activation of cAMP-dependent protein kinase, mediates many of the known effects of efferent input. A photoreceptor-specific protein, myosin III, is phosphorylated in response to efferent input; this protein may play a role in the efferent stimulated changes in photoreceptor structure and function. Anatomical, biophysical, biochemical, and molecular approaches are now being effectively combined in studies of Limulus eyes; thus, this preparation should be particularly useful for further detailed investigations of mechanisms underlying the modulation of primary sensory cells by efferent input.

Animals↗
PubMed2001

Photoreception and circadian clock system of the chicken pineal gland.

Chicken pinealocytes contain three major components of the circadian clock system: 1) a self-sustained oscillator, 2) a photic-input pathway to the oscillator, and 3) an overt output represented by the rhythmic production of melatonin. Even under cultured conditions of isolated pineal gland or dissociated pinealocytes, the input-oscillator-output functions are well maintained. Because of these experimental advantages, chicken pineal gland has been one of the best models for the study of the circadian clock system. Since the finding of a pineal-specific photoreceptive molecule, pinopsin, we have characterized the endogenous phototransduction pathway in the pinealocytes. On the other hand, despite the long history of chick pineal research, the molecular mechanism underlying the pineal clock oscillation has been largely unknown. Our recent characterization of the chick pineal clock genes strongly suggests that they constitute a transcription/translation-based autoregulatory feedback loop, which is very similar to that generating circadian rhythmicity in mammalian SCN.

Animals↗
PubMed2001

Molecular evidence that human ocular ciliary epithelium expresses components involved in phototransduction.

Here we report the expression, in the human ocular ciliary epithelium and in a human nonpigmented (NPE) ciliary epithelial cell line, of genes usually restricted to cone and rod photoreceptor cells of the retina. By RT-PCR and DNA sequencing we identified the expression of rhodopsin and components linked to its deactivation, including rhodopsin kinase, recoverin, and visual arrestin. We also detected the expression of transducin (T-alpha), phosphodiesterase (PDE-alpha), and cGMP-gated channel alpha-subunits. Cultured NPE cells responded to treatment with phorbol ester by enhancing the expression of rhodopsin mRNA three- to fourfold. Indirect immunofluorescence of the intact ciliary epithelium with monoclonal antibodies (MAbs) against rhodopsin, rhodopsin kinase, and visual arrestin revealed labeling preferentially restricted to the NPE cells. Furthermore, Western blot analysis of whole lysates from the pars plicata region of the human ciliary epithelium with MAbs demonstrated immunochemical cross-reactivity with proteins of molecular mass similar to rhodopsin (36 kDa), rhodopsin kinase (64 to 66 kDa), and arrestin (48-52 kDa) from the human retina. These results provide the first molecular evidence that components of a non-visual phototransduction pathway are expressed in the human ocular NPE ciliary epithelium, which may be linked to circadian entrainment tasks.

Animals↗
PubMed1999

Blue light signaling chains in Phycomyces: phototransduction of carotenogenesis and morphogenesis involves distinct protein kinase/phosphatase elements.

Carotenogenesis and morphogenesis represent two of the several responses sensitive to blue light which characterize the lower eukaryote Phycomyces blakesleeanus. Speculating that reversible phosphorylation may be an intracellular event beyond the photoperception step, we resorted to the use of first-choice inhibitors of protein phosphatases and protein kinases. The mycelial beta-carotene content of dark-grown cultures was induced by all agents administered, while the morphogenic output showed the typical trend effected by light only with one of the protein kinase inhibitors. Our data provide convincing evidence that protein phosphorylation plays a regulatory role in photocarotenogenesis and photomorphogenesis of Phycomyces. According to the model we propose, the putative signaling elements involved are anticipated to have a repressive function in the dark so that the responses are maintained in the "off" mode until the moment photon information has to flow through the regulatory circuit.

Culture Media↗
PubMed1999

Protein conformational changes in the bacteriorhodopsin photocycle.

We report a comprehensive electron crystallographic analysis of conformational changes in the photocycle of wild-type bacteriorhodopsin and in a variety of mutant proteins with kinetic defects in the photocycle. Specific intermediates that accumulate in the late stages of the photocycle of wild-type bacteriorhodopsin, the single mutants D38R, D96N, D96G, T46V, L93A and F219L, and the triple mutant D96G/F171C/F219L were trapped by freezing two-dimensional crystals in liquid ethane at varying times after illumination with a light flash. Electron diffraction patterns recorded from these crystals were used to construct projection difference Fourier maps at 3.5 A resolution to define light-driven changes in protein conformation. Our experiments demonstrate that in wild-type bacteriorhodopsin, a large protein conformational change occurs within approximately 1 ms after illumination. Analysis of structural changes in wild-type and mutant bacteriorhodopsins under conditions when either the M or the N intermediate is preferentially accumulated reveals that there are only small differences in structure between M and N intermediates trapped in the same protein. However, a considerably larger variation is observed when the same optical intermediate is trapped in different mutants. In some of the mutants, a partial conformational change is present even prior to illumination, with additional changes occurring upon illumination. Selected mutations, such as those in the D96G/F171C/F219L triple mutant, can sufficiently destabilize the wild-type structure to generate almost the full extent of the conformational change in the dark, with minimal additional light-induced changes. We conclude that the differences in structural changes observed in mutants that display long-lived M, N or O intermediates are best described as variations of one fundamental type of conformational change, rather than representing structural changes that are unique to the optical intermediate that is accumulated. Our observations thus support a simplified view of the photocycle of wild-type bacteriorhodopsin in which the structures of the initial state and the early intermediates (K, L and M1) are well approximated by one protein conformation, while the structures of the later intermediates (M2, N and O) are well approximated by the other protein conformation. We propose that in wild-type bacteriorhodopsin and in most mutants, this conformational change between the M1 and M2 states is likely to make an important contribution towards efficiently switching proton accessibility of the Schiff base from the extracellular side to the cytoplasmic side of the membrane.

Bacteriorhodopsins↗
PubMed2001

The molecular genetics of circadian rhythms in Arabidopsis.

While a number of physiological and biochemical processes in plants have been found to be regulated in a circadian manner, the mechanism underlying the circadian oscillator remains to be elucidated. Advances in the identification and characterization of components of the plant circadian system have been made largely through the use of genetics in Arabidopsis thaliana. Results so far indicate that the generation of rhythmicity by the Arabidopsis clock relies on molecular mechanisms that are similar to those described for other organisms, but that a totally different set of molecular components has been recruited to perform these functions.

Arabidopsis↗
PubMed2002

Calcium and phototransduction.

Visual phototransduction, the conversion of incoming light to an electrical signal, takes place in the outer segments of the rod and cone photoreceptor cells. Light reduces the concentration of cGMP, which, in darkness, keeps open cationic channels present in the plasma membrane of the outer segment. Ca2+ plays an important role in phototransduction by modulating the cGMP-gated channels as well as cGMP synthesis and breakdown. Ca2+ is involved in a negative feedback that is essential for photoreceptor adaptation to background illumination. The effects of Ca2+ on the different components of rod phototransduction have been characterized and can quantitatively account for the steady state responses of the rod cell to background illumination. The propagation of the Ca2+ feedback signal from the periphery toward the center of the outer segment depends on the Ca2+ diffusion coefficient, which has a value of 15 +/- 1 microm2 s(-1). This value shows that diffusion of Ca2+ in the radial direction is quite slow providing a significant barrier in the propagation of the feedback signal. Also, because the diffusion coefficient of Ca2+ is much smaller than that of cGMP, the decline of Ca2+ in the longitudinal direction lags behind the propagation of excitation by the decline of cGMP.

Animals↗
PubMed2004

The circadian Clock mutant mouse: impaired masking response to light.

Synchronization of an internal clock (entrainment) and a direct response to light (masking) are complementary ways of restricting activity of an animal to day or night. The protein CLOCK has an important role in the oscillatory mechanism of mammalian pacemakers. Our data show that it is also involved in masking responses. Mice with the Clock/Clock mutation reduced their wheel running less than wildtypes when given 1-h light pulses of light (2-1,600 lx) in the night. With dimmer lights (<2 lx), there were no significant differences between mutant and wildtype mice. Impaired masking responses to light in Clock/Clock mice were confirmed in tests with ultradian light-dark cycles (3.5:3.5 h and 1:1 h). Tests with pulses of light longer than 1 h revealed that, although the mutants responded more slowly to light, they sustained the suppression of activity over the course of the 3-h tests better than wildtypes.

Activity Cycles↗
PubMed2006

Of mice and women: light as a circadian stimulus in breast cancer research.

OBJECTIVE: Nocturnal rodents are frequently used as models in human breast cancer research, but these species have very different visual and circadian systems and, therefore, very different responses to optical radiation or, informally, light. Because of the impact of light on the circadian system and because recent evidence suggests that cancer risk might be related to circadian disruption, it is becoming increasingly clear that optical radiation must be properly characterized for both nocturnal rodents and diurnal humans to make significant progress in unraveling links between circadian disruption and breast cancer. In this paper, we propose a quantitative framework for comparing radiometric and photometric quantities in human and rodent studies. METHODS: We reviewed published research on light as a circadian stimulus for humans and rodents. Both suppression of nocturnal melatonin and phase shifting were examined as outcome measures for the circadian system. RESULTS: The data were used to develop quantitative comparisons regarding the absolute and spectral sensitivity for the circadian systems of humans and nocturnal rodents. CONCLUSIONS: Two models of circadian phototransduction, for mouse and humans, have been published providing spectral sensitivities for these two species. Despite some methodological variations among the studies reviewed, the circadian systems of nocturnal rodents are approximately 10,000 times more sensitive to optical radiation than that of humans. Circadian effectiveness of different sources for both humans and nocturnal rodents are offered together with a scale relating their absolute sensitivities. Instruments calibrated in terms of conventional photometric units (e.g., lux) will not accurately characterize the circadian stimulus for either humans or rodents.

Animals↗
PubMed2006

Light during darkness and cancer: relationships in circadian photoreception and tumor biology.

The relationship between circadian phototransduction and circadian-regulated processes is poorly understood. Melatonin, commonly a circadian phase marker, may play a direct role in a myriad of physiologic processes. The circadian rhythm for pineal melatonin secretion is regulated by the hypothalamic suprachiasmatic nucleus (SCN). Its neural source of light input is a unique subset of intrinsically photosensitive retinal ganglion cells expressing melanopsin, the primary circadian photopigment in rodents and primates. Action spectra of melatonin suppression by light have shown that light in the 446-477 nm range, distinct from the visual system's peak sensitivity, is optimal for stimulating the human circadian system. Breast cancer is the oncological disease entity whose relationship to circadian rhythm fluctuations has perhaps been most extensively studied. Empirical data has increasingly supported the hypothesis that higher risk of breast cancer in industrialized countries is partly due to increased exposure to light at night. Studies of tumor biology implicate melatonin as a potential mediator of this effect. Yet, causality between lifestyle factors and circadian tumor biology remains elusive and likely reflects significant variability with physiologic context. Continued rigorous empirical inquiry into the physiology and clinical implications of these habitual, integrated aspects of life is highly warranted at this time.

Animals↗
PubMed1994

Membrane association of phospholipase C encoded by the norpA gene of Drosophila melanogaster.

Severe mutations within the norpA gene of Drosophila abolish the photoreceptor potential and render the fly blind by deleting phospholipase C, an essential component of the phototransduction pathway. To study the membrane association of phospholipase C, we have utilized biochemical assays of phospholipase C activity, which predominant measurable phospholipase C activity in head homogenates has been shown to be encoded by norpA, as well as antisera generated against the major gene product of norpA to examine its subcellular distribution before and during phototransduction. We find that both phospholipase C activity and the norpA protein are predominantly associated with membrane fractions in heads of both light- and dark-adapted flies. Moreover, phospholipase C activity as well as norpA protein can be easily extracted from membrane preparations of light- or dark-adapted flies using high salt, indicating that the norpA protein is peripherally localized on the membrane. These data suggest that the norpA encoded phospholipase C of Drosophila is a permanent peripheral membrane protein. If this is indeed the case, then it would mean that the reversible redistribution of phospholipase C from the cytosol to the membrane, as observed in epidermal growth factor receptor stimulation of mammalian phospholipase C gamma, is not a universal mechanism utilized by all types of phosphatidylinositol-specific phospholipase C.

Animals↗
PubMed1995

Cloning and characterization of a G protein alpha-subunit-encoding gene from the basidiomycete, Coprinus congregatus.

Complementary DNA (cDNA) clones encoding two G protein alpha-subunit proteins (CGP alpha 1 and CGP alpha 2) were isolated from a Coprinus congregatus (Cc) hyphal tip cell (HTC) library using PCR-generated biotinylated G protein probes. Sequence analysis of the Cc cgp alpha 1 gene indicates that the gene contains an open reading frame (ORF) that translates into a putative 353-amino-acid (aa) product. The predicted CGP alpha 1 protein exhibits similarity to all known G protein alpha-subunits (it has all of the consensus regions for a GTP-binding protein), especially the mammalian retinal G protein, transducin. The CGP alpha 1 aa sequence is 50% identical overall to the transducin subfamily, cgp alpha 1 shares the same aa size grouping as transducin alpha-subunits and, unlike many other G proteins, both CGP alpha 1 and transducin seem to possess a cholera toxin (CTX)- and pertussis toxin (PTX)-sensitive site. Preliminary reverse transcription PCR (RT-PCR) analysis of cgp alpha 1 and cgp alpha 2 mRNA expression revealed that, unlike cgp alpha 2 which seems to be constitutively expressed, cgp alpha 1 is expressed only in HTC that are competent in responding to light. Thus, the cgp alpha 1 product, CGP alpha 1, is a likely candidate for regulating the blue light-induced signal transduction photomorphogenesis system found in Cc.

Amino Acid Sequence↗
PubMed1989

Chromaffin cell proliferation in the adult rat adrenal medulla.

Epinephrine and norepinephrine-containing chromaffin cells proliferate in the adrenal glands of normal adult rats throughout life. Moreover, their rate of proliferation is markedly increased by short-term administration of reserpine, one of many agents which in long-term experiments are associated with the development of adrenal medullary tumors. Current data suggest that chromaffin cell proliferation in the adult rat adrenal is mediated by the interaction of neurogenic and hormonal signals. Reserpine is known to directly deplete catecholamine stores, and to reflexively increase the activity of the splanchnic nerve endings innervating the adrenal medulla to stimulate both secretion and synthesis of catecholamines and other secretory granule constituents. Its effect on chromaffin cell proliferation suggests that the same signals may regulate chromaffin cell number to meet physiological needs. The reserpine model might shed light on signal transduction mechanisms which normally promote or prevent proliferation of chromaffin cells and of other neuroendocrine cells during development or in adult life, and on ways in which such mechanisms are altered in the course of the development and progression of tumors. It also suggests the possibility that chromaffin cells might be propagated in vitro for use in basic biological studies or in transplants for the treatment of Parkinson's disease.

Adrenal Medulla↗
PubMed2004

The phototransduction cascade in the isolated chick pineal gland revisited.

It is well established that the isolated chick pineal gland is directly light sensitive and that melatonin synthesis of the gland can be inhibited by exposing the gland to light during scotophase. Since not all the steps of the phototransduction cascade have been clarified to the same extent as in the retina, we have treated isolated chick pineal glands with 90 min of light during scotophase and with drugs that affect key-components of vertebrate phototransduction, i.e., cyclic guanosine monophosphate (cGMP) phosphodiesterase 6 (PDE6), cGMP levels and cGMP-gated calcium channels. The endpoint measured was the activity of the rate-limiting enzyme of melatonin synthesis, arylalkylamine N-acetyltransferase (AA-NAT), which is inhibited by light. The effects on AA-NAT activity of light were negated by addition of dipyridamol and zaprinast, either of which inhibits the light-induced activation of PDE6. The effect of light was also counteracted by the nitric oxide donor sodium nitroprusside and C-type natriuretic peptide, both of which increase cGMP levels, and by the calcium channel agonist Bay K 8644, which prevents the cGMP-decrease-induced closure of cGMP-gated calcium channels. Inhibition of nitric oxide synthase (NOS) by N(G)-nitro-l-arginine did not influence the inhibitory effect of light, suggesting that the NOS pathway does not play a role. Since the light effect on AA-NAT activity involves both cGMP and cyclic adenosine monophosphate (cAMP) hydrolysis, we have also studied whether the cGMP-inhibited cAMP phosphodiesterase 3 (PDE3) is involved. As the specific PDE3 inhibitor cilostamide is without effect, we assume that the light-induced decrease of cAMP levels does not involve PDE3. These results taken together strongly suggest that the investigated steps of the phototransduction cascade in the isolated chick pineal gland are basically similar to those in the retina.

3',5'-Cyclic-AMP Phosphodiesterases↗
PubMed2004

Circadian and cAMP-dependent modulation of retinal cone cGMP-gated channels does not require protein synthesis or calcium influx through L-type channels.

Circadian oscillators of chicken retinal cone photoreceptors modulate the gating properties of cGMP-gated channels (CNGCs) such that they have a higher apparent affinity for cGMP during the subjective night. This effect is driven in part by cAMP, which acts through Erk MAP kinase to initiate a cascade leading to modulation of CGNCs. Here, we show that cAMP effects on the gating properties CNGCs persist when protein synthesis is blocked. The effects is cAMP also persist when calcium influx through L-type channels is blocked by nitrendipine. The mechanisms whereby cAMP modulates CNGCs therefore differ from those previously reported to underline regulation of melatonin synthesis and secretion.

Animals↗