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The Journal of Neuroscience, January 15, 2000, 20(2):600-605
A Novel Human Opsin in the Inner Retina
Ignacio
Provencio1,
Ignacio R.
Rodriguez2,
Guisen
Jiang1,
William
Pär
Hayes3,
Ernesto F.
Moreira2, and
Mark D.
Rollag1
1 Department of Anatomy and Cell Biology, Uniformed
Services University of the Health Sciences, Bethesda, Maryland 20814, 2 National Eye Institute, National Institutes of Health,
Bethesda, Maryland 20892, and 3 Department of Biology, The
Catholic University of America, Washington, DC 20064
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ABSTRACT |
Here we report the identification of a novel human opsin,
melanopsin, that is expressed in cells of the mammalian inner retina. The human melanopsin gene consists of 10 exons and is mapped to chromosome 10q22. This chromosomal localization and gene structure differs significantly from that of other human opsins that typically have four to seven exons. A survey of 26 anatomical sites indicates that, in humans, melanopsin is expressed only in the eye. In
situ hybridization histochemistry shows that melanopsin
expression is restricted to cells within the ganglion and amacrine cell
layers of the primate and murine retinas. Notably, expression is not observed in retinal photoreceptor cells, the opsin-containing cells of
the outer retina that initiate vision. The unique inner retinal
localization of melanopsin suggests that it is not involved in image
formation but rather may mediate nonvisual photoreceptive tasks, such
as the regulation of circadian rhythms and the acute suppression of
pineal melatonin. The anatomical distribution of melanopsin-positive
retinal cells is similar to the pattern of cells known to project from
the retina to the suprachiasmatic nuclei of the hypothalamus, a primary
circadian pacemaker.
Key words:
circadian; melanopsin; opsin; photoreceptor; retina; retinal ganglion cell
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INTRODUCTION |
Light is a potent regulator of many
physiological processes in vertebrates. For example, the
synchronization of circadian rhythms to the 24 hr solar cycle and the
proper seasonal timing of reproduction are greatly influenced by
environmental light cues. Among nonmammalian vertebrates, many of these
processes persist in the absence of eyes (Groos, 1982 ). This finding
indicates that some effects of light are mediated through extraocular
photoreceptors. Several extraocular tissues in nonmammalian vertebrates
have been shown to be directly photosensitive in culture. These include the pineal gland (Deguchi, 1979), the iris (Barr and Alpern, 1963), and
dermal melanophores (Seldenrijk et al., 1979). Within the past 5 years,
multiple novel opsins have been identified and localized to extraocular
sites in nonmammalian vertebrates. Pinopsin is found in the pineal
gland of birds (Okano et al., 1994; Max et al., 1995; Kawamura and
Yokoyama, 1996) and reptiles (Kawamura and Yokoyama, 1997) and in the
brain of toads (Yoshikawa et al., 1998). Parapinopsin is present in the
parapineal gland of the catfish (Blackshaw and Snyder, 1999).
Melanopsin is localized to various extraretinal tissues in amphibians,
including dermal melanophores, deep brain nuclei, and iris (Provencio
et al., 1998b). Melanopsin and vertebrate ancient opsin have also been
observed in nonphotoreceptor cells of the inner retina of frogs and
fish, respectively (Provencio et al., 1998b; Soni et al., 1998).
Mammals also detect light for the entrainment of circadian rhythms and
the regulation of pineal melatonin production. In contrast to
nonmammalian vertebrates, the photoreceptors mediating nonvisual photic
processes are ocular in mammals (Nelson and Zucker, 1981). However, the
rod and cone photoreceptors of the retina are not required for the
regulation of circadian rhythms or acute suppression of pineal
melatonin levels (Freedman et al., 1999; Lucas et al., 1999). The
ocular photoreceptors mediating these nonvisual tasks in mammals remain
unknown. Having localized melanopsin to amphibian tissues known to
contain extraretinal photoreceptors and to nonphotoreceptor cells of
the amphibian retina (Provencio et al., 1998b), we hypothesize that
melanopsin expression can be used as a tool to identify candidate nonvisual photoreceptors in other vertebrates, including mammals.
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MATERIALS AND METHODS |
Cloning of human melanopsin. A lambda human
genomic DNA library (Clontech, Palo Alto, CA) was screened with a 475 bp oligonucleotide probe derived from the chicken melanopsin cDNA
sequence (M. D. Rollag, I. Provencio, and M. Zatz, unpublished data).
DNA from a positive partial clone was restriction mapped, subcloned
into pGEM3Zf(+) (Promega, Madison, WI), and sequenced (PE Biosystems, Foster City, CA). A gene-specific primer pair was derived from the
sequence of the partial genomic clone (primer IPF85,
5'-GGAGGAGAGAAGGCACACAG-3'; primer IPB83, 5'-GCTGCTGCAG-ATGTCACAAT-3').
These primers were used to screen for a Genome Systems (St. Louis, MO)
PAC clone (number 20775) containing the entire melanopsin gene. The
Coriell Monochromosomal Panel 2 (Coriell Cell Repositories, Camden, NJ) was PCR screened with primers IPF85 and IPB83 to establish a gross chromosomal localization. A finer localization was achieved by a PCR
screen with the same primers against the Stanford Human Genome Center
G3 radiation hybrid panel (Stewart et al., 1997). PCR primers were
also designed within putative coding regions of the genomic
sequence as determined by homology to Xenopus melanopsin (primer IPF70, 5'-GACACCCTACATGAGCTCGG-3'; primer IPB64,
5'-CTGTACTTGGGGTGGGTGAT-3'). These primers were used to amplify a 95 bp
melanopsin cDNA fragment by PCR (PE Biosystems) from human
retina cDNA. A full-length cDNA was obtained by 5' and 3' rapid
amplification of cDNA ends (RACE) (Life Technologies, Rockville, MD).
Reverse transcription-PCR of multiple tissues. Human total
RNA was obtained (Clontech) or isolated from tissues (National Disease
Research Interchange, Philadelphia, PA) by the guanidinium thiocyanate
method (Chomczynski and Sacchi, 1987) and reverse transcribed
onto magnetic beads as described previously (Rodriguez et al., 1994).
One microliter of solid phase cDNA from each tissue was subjected to
PCR amplification (AmpliTaq Gold polymerase; PE Biosystems) for 20 cycles using human glyceraldehyde-3-phosphate dehydrogenase
(GAPDH)-specific primers (primer GAPDH forward, 5'-CCACCCATGGCAAATTCCATGGCA-3'; primer GAPDH reverse,
5'-TCTAGACGGCAGGTCAGGTCCACC-3') or 30 cycles using human
melanopsin-specific primers (primer IPF108, 5'-ACTCAGGATGAACCCTCCTTC-3'; primer IPB91,
5'-TGAACATGTTGGCAGGTGTC-3'). The GAPDH and melanopsin PCR reactions
from each tissue were mixed 1:1 (v/v), and 5 µl of each mixture was
electrophoresed and visualized by ethidium bromide staining. The
expected products for the melanopsin and GAPDH primer pairs from cDNA
template are 334 and 600 bp, respectively. The expected products for
the melanopsin and GAPDH primer pairs from genomic DNA template are
3292 and 1104 bp, respectively.
Cloning of mouse melanopsin. A 228 bp mouse (Mus
musculus) melanopsin cDNA fragment was PCR-amplified (AmpliTaq
Gold polymerase; PE Biosystems) from mouse retina cDNA using human
melanopsin-specific primers (primer IPF82, 5'-ATCCTGCTCCTGGGACTACA-3';
primer IPB74, 5'-ATCTTGGCCATCTTGCACTC-3'). A full-length cDNA was
subsequently obtained by 5' and 3' RACE (Life Technologies).
In situ hybridization histochemistry. Adult macaque
(Macaca mulatta) and adult and juvenile (postnatal day 10)
mouse eyes were removed. The corneas and lenses were dissected away
from the adult eyes. The resulting tissues were either immersion fixed (4% formalin, 24 hr), frozen in Tissue-Tek (Miles, Elkhart, IN) and
immediately freeze-mounted, or placed into TRI-reagent (Sigma, St. Louis, MO) for RNA isolation. Total RNA was isolated by the guanidinium thiocyanate method (Chomczynski and Sacchi, 1987). A 417 bp
cDNA fragment of the macaque melanopsin homolog was obtained by reverse
transcription (RT)-PCR using primers designed from the
human melanopsin cDNA sequence (primer IPF82,
5'-ATCCTGCTCCTGGGACTACA-3'; primer IPB64,
5'-CTGTACTTGGGGTGG- GTGAT-3'). Products were cloned into pCRII-TOPO
(Invitrogen, Carlsbad, CA) and sequenced. A 957 bp mouse melanopsin
cDNA fragment was similarly cloned using mouse melanopsin-specific
primers (primer IPF107, 5'-TCTTCATCTTCAGGGCCATC-3'; primer IPB116,
5'-TTCTCTGCTGTAGGCCACATA-3'). Plasmids were used to generate
35S-labeled antisense and sense control
riboprobes (Ambion, Austin, TX). Eyecups were sectioned (15 µm),
thaw-mounted onto silanized RNase-free microscope slides, and
hybridized with the riboprobes as described previously (Shi and Hayes,
1994).
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RESULTS |
Cloning of human melanopsin
We determined the nucleotide sequence of both strands of the human
melanopsin gene, including exons, introns, and 2.6 kb of the 5'
flanking region (GenBank accession number AF147788). All introns are
flanked by consensus splice sites. The melanopsin gene has 10 exons
distributed over 11.8 kb. Its gene structure is unique among the
vertebrate opsins as evidenced by intron positions significantly
different from the intron positions of the rod and cone opsins (Fig.
1). PCR screens of the radiation hybrid
panels mapped the melanopsin gene to human chromosome 10q22. A 2.3 kb melanopsin cDNA containing a 1.4 kb open reading frame (ORF) was cloned from human retina cDNA, indicating that melanopsin is normally transcribed in the human eye. RT-PCR from 26 human anatomical sites
demonstrated melanopsin expression in the eyes but in none of the other
sites examined (Fig. 2). Unlike
Xenopus, human melanopsin expression was not observed in the
pineal gland or any brain region examined.
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Figure 1.
Melanopsin differs from other human opsins.
A, Structure of melanopsin gene. The 5' and 3'
untranslated regions of exons 1 and 10, respectively, are indicated in
white. B, Comparison of human melanopsin,
rhodopsin (Nathans and Hogness, 1984), blue cone opsin (Nathans et al.,
1986), red and green cone opsins (Nathans et al., 1986), RGR (Shen et
al., 1994), peropsin (Hui et al., 1997), and encephalopsin (Blackshaw
and Snyder, 1999) ORFs. Portions of the ORFs corresponding to
the transmembrane domains are shown in white and are
labeled. Positions of introns are indicated ( ), and the respective
chromosomal locations are displayed to the right.
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Figure 2.
Human melanopsin is expressed in the eye. RT-PCR
from multiple human tissues of melanopsin (334 bp) and the GAPDH
positive control (600 bp). The faint melanopsin product from
RPE/choroid may have resulted from retinal contamination during
dissection. Melanopsin is not expressed in the other tissues
examined.
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Cloning of mouse melanopsin
We cloned a 2.1 kb melanopsin cDNA with a 1.6 kb open reading
frame from mouse retina cDNA (GenBank accession number AF147789). When
the human and mouse cDNAs are translated and aligned, the predicted
transmembrane and loop domains are 86% identical to each other (Fig.
3) and 57% identical to the frog
homolog. The cytoplasmic and extracellular tails are significantly
different in sequence and length.
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Figure 3.
Alignment of human and mouse melanopsin-deduced
amino acid sequences. Sequences were aligned with ClustalW 1.6 (Thompson et al., 1994). Predicted transmembrane domains are
boxed and were determined by homology to
Xenopus melanopsin (Provencio et al., 1998b). The
Schiff's base lysine ( ) and the invertebrate-like tyrosine
counterion ( ) are indicated.
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In situ hybridization histochemistry
In primates, a low level of melanopsin expression is observed in
many retinal cells within the ganglion cell layer of the retina (Fig.
4). Higher levels of message are found in
sparsely distributed cells within the inner lamina of the inner nuclear layer among amacrine cell perikarya. In the mouse retina, melanopsin is
expressed in only a few cells in the ganglion cell layer and even fewer
cells in the amacrine cell layer (Fig.
5).
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Figure 4.
Melanopsin is expressed in the monkey inner
retina. Bright-field (A) and dark-field
(B) photomicrographs of a section of monkey
retina probed with an antisense monkey melanopsin riboprobe.
C, An adjacent section probed with a sense control
riboprobe. GC, Ganglion cell layer; INL,
inner nuclear layer; OD, optic disk; P,
photoreceptor layer. Scale bar, 150 µm.
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Figure 5.
Melanopsin is expressed in the mouse inner retina.
A, Cross-section of a 10-d-old mouse eye probed with an
antisense mouse melanopsin riboprobe. B,
C, Bright-field and dark-field photomicrographs of
indicated cell within the amacrine cell layer in A.
D, E, Bright-field and dark-field
photomicrographs of indicated cell pair within the ganglion cell layer
in A. GC, Ganglion cell layer;
INL, inner nuclear layer; P,
photoreceptor layer. Scale bars: A, 250 µm;
B, 50 µm.
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DISCUSSION |
Perhaps the most striking feature that distinguishes melanopsin
from other human opsins is its greater sequence homology with the
invertebrate opsins than those of vertebrates. In fact, the predicted
amino acid sequence is more similar to the
Gq-coupled opsin of the scallop (Kojima et al.,
1997) than to any known vertebrate opsin. The similarity to the
invertebrate photopigments is also apparent in several predicted
structural and biochemical features. This includes the substitution of
the acidic Schiff's base "counterion," typical of
vertebrate opsins, with an aromatic residue that is typical of the
invertebrate opsins. The absence of this acidic counterion in
melanopsin suggests that the photopigment-regenerating mechanism more
closely resembles that of the invertebrates (Gärtner and Towner,
1995). Invertebrate opsin-based photopigments retain their
retinaldehyde chromophore after it is photoisomerized from the
11-cis to all-trans configuration. The retained
chromophore is reisomerized to the 11-cis configuration by a
second wavelength of light. This in situ photopigment
regeneration does not require proximity to an auxiliary
chromophore-regenerating tissue, such as the retinal pigment epithelium
(RPE). Such anatomical independence is a valuable attribute for
nonvisual photoreceptors that may reside in a wide variety of tissues.
Indeed, cells containing melanopsin transcripts in the retina are not
juxtaposed to the RPE but rather are situated within neural elements of
the inner retina.
The inner retinal distribution of melanopsin-positive cells shares a
remarkable resemblance to the cohort of retinal cells known to project
to the primary circadian pacemaker of rodents, the suprachiasmatic
nuclei (SCN) of the hypothalamus (Pickard, 1980, 1982; Moore et al.,
1995). In mice, only a small subset of widely distributed retinal
ganglion and amacrine-like cells project to the SCN (Balkema and
Dräger, 1990; Provencio et al., 1998a). The number and location
of these cells are similar to that of the melanopsin-positive cells.
The presence of melanopsin in the inner retina raises the
possibility that some mammalian ganglion and amacrine cells are directly photosensitive. This possibility is consistent with the finding that naturally occurring and transgenic mice that lack rods and
cones, although maintaining an apparently normal inner retina, are
capable of photoregulating circadian locomotor activity rhythms and
pineal melatonin levels in a manner indistinguishable from wild-type
controls (Foster et al., 1991; Provencio et al., 1994; Freedman et al.,
1999). Bilateral removal of the eyes abolishes such regulation (Nelson
and Zucker, 1981). Together, these data indicate that, whereas the eyes
are required for the effect of light on the circadian axis, the visual
photoreceptors are not necessary. This paradox strongly suggests that
some class of nonrod, noncone photoreceptor exists within the mammalian
eye. Ocular nonvisual photoreception would explain why some humans
retain an ability to acutely suppress serum melatonin concentrations in
response to light exposure despite being cognitively and clinically blind (Czeisler et al., 1995).
The prospect that retinal ganglion cells are photoreceptive has
received much attention because of the discovery that many mouse
ganglion cells contain cryptochromes, a class of blue light-absorbing, flavin-based photopigments related to DNA photolyases (Miyamoto and
Sancar, 1998). Two cryptochromes (mCRY1 and mCRY2) have been localized
to ganglion and inner nuclear cells and have been proposed as candidate
circadian photopigments. Knock-out mice missing one or both
cryptochromes have been constructed to assess their role in circadian
rhythm regulation (Thresher et al., 1998; van der Horst et al., 1999).
One would expect the phenotype of a circadian photopigment-deficient
mouse to be like that of a bilaterally enucleated mouse (Nelson and
Zucker, 1981). That is to say, a mouse incapable of circadian
photoreception should be unable to entrain its circadian locomotor
activity rhythms to the light/dark cycle. Instead, it has been found
that mice lacking either of the cryptochromes remain capable of
photoentrainment (Thresher et al., 1998; van der Horst et al., 1999;
Vitaterna et al., 1999). Mice missing both cryptochromes are
arrhythmic, probably as a result of a nonfunctioning clock. This latter
finding makes the analysis of entrainment in cryptochrome double
knock-out mice difficult. Thus, the issue of whether mice lacking both
cryptochromes can photoentrain remains unresolved (van der Horst et
al., 1999; Vitaterna et al., 1999). Together, these data from knock-out
mice fail to show that cryptochromes act as circadian photopigments. Rather, they suggest that cryptochromes are critical components of the
molecular machinery of the clock. This conclusion has been strengthened
by investigators who have shown that cryptochromes are indeed essential
components of the mammalian circadian clock (Griffin et al., 1999; Kume
et al., 1999). Furthermore, there is evidence that the functional
interaction of mammalian cryptochromes with other clock proteins is
independent of light, again strongly suggesting that cryptochromes are
not acting as circadian photopigments within the circadian system of
mammals (Griffin et al., 1999).
An independent method to discriminate whether flavin- or opsin-based
photopigments mediate particular light responses is through action
spectrum analysis (Lucas and Foster, 1999). Action spectra of rodent
circadian responses more closely resemble the spectral absorbance
profile of opsin-based photopigments rather than flavin-based cryptochromes (Takahashi et al., 1984; Provencio and Foster, 1995; Yoshimura and Ebihara, 1996). Melanopsin is one of four opsins expressed outside the photoreceptor layer in human and mouse retinas, the other three being retinal G-protein-coupled receptor (RGR), peropsin, and encephalopsin. RGR is found predominantly in
intracellular membranes of the RPE and Müller cells (Pandey et
al., 1994) and functions as a photoisomerase (Hao and Fong, 1999).
Peropsin is found within the microvilli of RPE cells, and its function
is unknown (Hui et al., 1997). The function of encephalopsin is also unknown, and it has not yet been shown to be expressed in the eye
(Blackshaw and Snyder, 1999). Melanopsin is the only one of the four
known mammalian nonvisual opsins that is expressed by cells in the
ganglion and amacrine cell layers of the retina. This unique anatomical
localization, coupled with the known action spectra for mammalian
circadian photoregulation, makes melanopsin a viable candidate as a
mammalian circadian photopigment.
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FOOTNOTES |
Received Aug. 16, 1999; revised Nov. 2, 1999; accepted Nov. 4, 1999.
This work was supported by National Science Foundation Grant
IBN-9809916 and Uniformed Services University of the Health Sciences Grant RO7049 (both to M.D.R.). We thank Dr. Paul Russell for the monkey
eye tissue, Dr. Diane Borst for the mouse tissue, and Dr. Michael Flora
for helpful discussions.
Correspondence should be addressed to Mark D. Rollag, Department of
Anatomy, Uniformed Services University of the Health Sciences, 4301 Jones Bridge Road, Bethesda, MD 20814. E-mail: mrollag{at}usuhs.mil.
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C. L. Thompson, W. S. Blaner, R. N. Van Gelder, K. Lai, L. Quadro, V. Colantuoni, M. E. Gottesman, and A. Sancar
Preservation of light signaling to the suprachiasmatic nucleus in vitamin A-deficient mice
PNAS,
September 25, 2001;
98(20):
11708 - 11713.
[Abstract]
[Full Text]
[PDF]
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C. P. Selby, C. Thompson, T. M. Schmitz, R. N. Van Gelder, and A. Sancar
Functional redundancy of cryptochromes and classical photoreceptors for nonvisual ocular photoreception in mice
PNAS,
December 19, 2000;
97(26):
14697 - 14702.
[Abstract]
[Full Text]
[PDF]
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J. Hannibal, P. Hindersson, S. M. Knudsen, B. Georg, and J. Fahrenkrug
The Photopigment Melanopsin Is Exclusively Present in Pituitary Adenylate Cyclase-Activating Polypeptide-Containing Retinal Ganglion Cells of the Retinohypothalamic Tract
J. Neurosci.,
January 1, 2002;
22(1):
RC191 - RC191.
[Abstract]
[Full Text]
[PDF]
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ABSTRACT
INTRODUCTION
Gene
