122:. Based on action spectroscopy and simultaneous recordings of photocurrents and flagellar beating, it was determined that the photoreceptor currents and subsequent flagellar movements are mediated by rhodopsin and control phototaxis and photophobic responses. The extremely fast rise of the photoreceptor current after a brief light flash led to the conclusion that the rhodopsin and the channel are intimately linked in a protein complex, or even within one single protein.
399:'s lab optimized ChR2 for further increases in steady-state conductance and dramatically reduced desensitization by creating chimeras of ChR1 and ChR2 and mutating specific amino acids, yielding ChEF and ChIEF, which allowed the driving of trains of action potentials up to 100 Hz. In 2010, the groups of Hegemann and Deisseroth introduced an E123T mutation into native ChR2, yielding ChETA, which has faster on- and off-
375:, deploying the indirectly light-gated ion channel P2X2, it was henceforth microbial opsins like channelrhodopsin that dominated the field of genetically targeted remote control of excitable cells, due to the power, speed, targetability, ease of use, and temporal precision of direct optical activation, not requiring any external chemical compound such as caged ligands.
5157:
520:. Searches for homologous sequences in other organisms has yielded spectrally improved and stronger red-shifted channelrhodpsins (Chrimson). In combination with ChR2, these yellow/red light-sensitive channelrhodopsins allow controlling two populations of neurons independently with light pulses of different colors.
498:
Mutating the E123 residue accelerates channel kinetics (ChETA), and the resulting ChR2 mutants have been used to spike neurons at up to 200 Hz. In general, channelrhodopsins with slow kinetics are more light-sensitive on the population level, as open channels accumulate over time even at low light levels.
97:
and photoorientation of microalgae have been studied over more than a hundred years in many laboratories worldwide. In 1980, Ken Foster developed the first consistent theory about the functionality of algal eyes. He also analyzed published action spectra and complemented blind cells with retinal and
687:
Neurons can tolerate ChR expression for a long time, and several laboratories are testing optogenetic stimulation to solve medical needs. In blind mice, visual function can be partially restored by expressing ChR2 in inner retinal cells. In 2021, the red-light sensitive ChR ChrimsonR was virally
506:
H134R and T159C mutants display increased photocurrents, and a combination of T159 and E123 (ET/TC) has slightly larger photocurrents and slightly faster kinetics than wild-type ChR2. ChIEF, a chimera and point mutant of ChR1 and ChR2, demonstrates large photocurrents, little desensitization and
497:
Closing of the channel after optical activation can be substantially delayed by mutating the protein residues C128 or D156. This modification results in super-sensitive channelrhodopsins that can be opened by a blue light pulse and closed by a green or yellow light pulse (Step-function opsins).
515:
Chimeric channelrhodopsins have been developed by combining transmembrane helices from ChR1 and VChR1, leading to the development of ChRs with red spectral shifts (such as C1V1 and ReaChR). ReaChR has improved membrane trafficking and strong expression in mammalian cells, and has been used for
410:
The groups of
Hegemann and Deisseroth also discovered that the introduction of the point mutation C128S makes the resulting ChR2-derivative a step-function tool: Once "switched on" by blue light, ChR2(C128S) stays in the open state until it is switched off by yellow light – a modification that
353:
as the light-sensing co-factor and it was unclear whether central mammalian nerve cells would contain sufficient retinal levels, but they do. It also showed, despite the small single-channel conductance, sufficient potency to drive mammalian neurons above action potential threshold. From this,
184:
550:
Mutating E90 to the positively charged amino acid arginine turns channelrhodopsin from an unspecific cation channel into a chloride-conducting channel (ChloC). The selectivity for Cl- was further improved by replacing negatively charged residues in the channel pore, making the
527:. After some engineering to improve membrane trafficking and speed, the resulting tool (CheRiff) produced large photocurrents at 460 nm excitation. It has been combined with the Genetically Encoded Calcium Indicator jRCaMP1b in an all-optical system called the OptoCaMP.
308:(ACRs) and potassium-selective channelrhodpsins (HcKCR1, HcKCR2) were structurally analyzed to understand their ion selectivity. ACRs and KCRs have been used to inhibit neuronal activity. Recently discovered viral channelrhodopsins (VCR1) are localized to the membrane of the
680:
can be stimulated to perform some desired behaviors for applications in robotics and control. ChR2 has also been used to map long-range connections from one side of the brain to the other, and to map the spatial location of inputs on the dendritic tree of individual neurons.
415:. VChR1 produces only tiny photocurrents, but with an absorption spectrum that is red-shifted relative to ChR2. Using parts of the ChR1 sequence, photocurrent amplitude was later improved to allow excitation of two neuronal populations at two distinct wavelengths.
448:
In March 2013, the Brain Prize (Grete
Lundbeck European Brain Research Prize) was jointly awarded to Bamberg, Boyden, Deisseroth, Hegemann, Miesenböck, and Nagel for "their invention and refinement of optogenetics". The same year, Hegemann and Nagel received the
354:
channelrhodopsin became the first optogenetic tool, with which neural activity could be controlled with the temporal precision at which neurons operate (milliseconds). A second study was published later confirming the ability of ChR2 to control the activity of
371:. This was the first using ChR2 to steer the behavior of an animal in an optogenetic experiment, rendering a genetically specified cell type subject to optical remote control. Although both aspects had been illustrated earlier that year by the group of
125:
The name "channelrhodopsin" was coined to highlight this unusual property, and the sequences were renamed accordingly. The nucleotide sequences of the rhodopsins now called channelrhodopsins ChR1 and ChR2 were finally uncovered in a large-scale
435:(opto-fMRI). Other labs have pioneered the combination of ChR2 stimulation with calcium imaging for all-optical experiments, mapping of long-range and local neural circuits, ChR2 expression from a transgenic locus – directly or in the
390:
H134R (exchanging the amino acid
Histidine in position 134 of the native protein for an Arginine) resulted in increased steady-state conductance, as described in a 2005 paper that also established ChR2 as an optogenetic tool in
563:
ACR) inhibit neuronal spiking in cell culture and in intact animals when illuminated with blue light. Calcium-selective channelrhodopsins have been engineered to activate calcium-dependent enzymes in cells.
365:
It was demonstrated that ChR2, if expressed in specific neurons or muscle cells, can evoke predictable behaviors, i.e. can control the nervous system of an intact animal, in this case the invertebrate
4613:
Lagali PS, Balya D, Awatramani GB, Münch TA, Kim DS, Busskamp V, et al. (June 2008). "Light-activated channels targeted to ON bipolar cells restore visual function in retinal degeneration".
883:
Foster KW, Saranak J, Patel N, Zarilli G, Okabe M, Kline T, et al. (October 1984). "A rhodopsin is the functional photoreceptor for phototaxis in the unicellular eukaryote
Chlamydomonas".
485:
can be useful to visualize the morphology of ChR2 expressing cells, i.e. simultaneously indicate which cells are tagged with FP and allow the activity to be controlled by the channelrhodopsin.
362:-4 in this case, demonstrating for the first time that excitable cells could be activated and silenced using these two tools simultaneously, illuminating the tissue at different wavelengths.
539:(excitatory). Variants with moderate to high calcium permeability have been engineered (CatCh, CapChRs). K-specific channelrhodopsins (KCRs, WiChR) were recently discovered in various
1155:
Suzuki T, Yamasaki K, Fujita S, Oda K, Iseki M, Yoshida K, et al. (February 2003). "Archaeal-type rhodopsins in
Chlamydomonas: model structure and intracellular localization".
175:
Their roles in generation of photoreceptor currents in algal cells were characterized by Oleg
Sineshchekov, Kwang-Hwan Jung and John Spudich, and Peter Berthold and Peter Hegemann.
507:
kinetics similar to wild-type ChR2. Variants with extended open time (ChR2-XXL) produce extremely large photocurrents and are very light sensitive on the population level.
696:
have been shown to work well in animal experiments and are currently undergoing clinical trials. In the future, ChRs may find even more medical applications, e.g. for
300:Å. Within milliseconds, the retinal relaxes back to the all-trans form, closing the pore and stopping the flow of ions. Most natural channelrhodopsins are nonspecific
411:
deteriorates temporal precision, but increases light sensitivity by two orders of magnitude. They also discovered and characterized VChR1 in the multicellular algae
2729:"In-depth activation of channelrhodopsin 2-sensitized excitable cells with high spatial resolution using two-photon excitation with a near-infrared laser microbeam"
1360:
Bamann C, Kirsch T, Nagel G, Bamberg E (January 2008). "Spectral characteristics of the photocycle of channelrhodopsin-2 and its implication for channel function".
378:
To overcome its principal downsides – the small single-channel conductance (especially in steady-state), the limitation to one optimal excitation wavelength (~470
661:
ACR2 and the red-light sensitive cation channel
Chrimson which have been combined in a single protein (BiPOLES) for bidirectional control of membrane potential.
489:
close to the retinal binding pocket have been shown to affect the biophysical properties of the channelrhodopsin, resulting in a variety of different tools.
349:, achieving temporal precision on the order of milliseconds (both in terms of delay to spiking and in terms of temporal jitter). Because all opsins require
727:
Nagel G, Ollig D, Fuhrmann M, Kateriya S, Musti AM, Bamberg E, et al. (June 2002). "Channelrhodopsin-1: a light-gated proton channel in green algae".
74:
are the first discovered channelrhodopsins. Variants that are sensitive to different colors of light or selective for specific ions (ACRs, KCRs) have been
657:
together enable multiple-color optical activation and silencing of neural activity. Another interesting pair is the blue-light sensitive chloride channel
1675:
Boyden ES, Zhang F, Bamberg E, Nagel G, Deisseroth K (September 2005). "Millisecond-timescale, genetically targeted optical control of neural activity".
934:
Litvin FF, Sineshchekov OA, Sineshchekov VA (February 1978). "Photoreceptor electric potential in the phototaxis of the alga
Haematococcus pluvialis".
3577:
Govorunova EG, Gou Y, Sineshchekov OA, Li H, Wang Y, Brown LS, et al. (2021-09-17). "Kalium rhodopsins: Natural light-gated potassium channels".
5121:
382:
nm, blue) as well as the relatively long recovery time, not permitting controlled firing of neurons above 20–40 Hz – ChR2 has been optimized using
4085:
Zhang F, Wang LP, Brauner M, Liewald JF, Kay K, Watzke N, et al. (April 2007). "Multimodal fast optical interrogation of neural circuitry".
5047:
358:
neurons, at this time in the chick spinal cord. This study was the first wherein ChR2 was expressed alongside an optical silencer, vertebrate
5116:
4434:
5188:
2573:
Petreanu L, Huber D, Sobczyk A, Svoboda K (May 2007). "Channelrhodopsin-2-assisted circuit mapping of long-range callosal projections".
454:
5111:
642:
have been engineered and were also found in nature. These tools can be used to silence neurons in cell culture and in live animals by
256:). This makes cellular depolarization extremely fast, robust, and useful for bioengineering and neuroscience applications, including
5074:
1720:"Fast noninvasive activation and inhibition of neural and network activity by vertebrate rhodopsin and green algae channelrhodopsin"
1142:
639:
556:
305:
4407:
Xu Z, Ziye X, Craig H, Silvia F (Dec 2013). "Spike-based indirect training of a spiking neural network-controlled virtual insect".
269:
1141:
Kateriya, S. Fuhrmann, M. Hegemann, P.: Direct
Submission: Chlamydomonas reinhardtii retinal binding protein (cop4) gene; GenBank
450:
442:
535:
Most channelrhodopsins are unspecific cation channels. When expressed in neurons, they conduct mostly Na ions and are therefore
4028:"Multiple-color optical activation, silencing, and desynchronization of neural activity, with single-spike temporal resolution"
1256:"Channelrhodopsin-1 initiates phototaxis and photophobic responses in chlamydomonas by immediate light-induced depolarization"
608:
neurons without adding any chemical compounds. Before the discovery of channelrhodopsins, neuroscientists were limited to
1782:"Light activation of channelrhodopsin-2 in excitable cells of Caenorhabditis elegans triggers rapid behavioral responses"
1870:
Zhang F, Wang LP, Boyden ES, Deisseroth K (October 2006). "Channelrhodopsin-2 and optical control of excitable cells".
3610:"WiChR, a highly potassium-selective channelrhodopsin for low-light one- and two-photon inhibition of excitable cells"
1570:
544:
204:
162:
by
Takahashi's group. Both sequences were found to function as single-component light-activated cation channels in a
4951:"Ectopic expression of a microbial-type rhodopsin restores visual responses in mice with photoreceptor degeneration"
4566:"Ectopic expression of a microbial-type rhodopsin restores visual responses in mice with photoreceptor degeneration"
4517:"Ectopic expression of a microbial-type rhodopsin restores visual responses in mice with photoreceptor degeneration"
5040:
2027:
Gunaydin LA, Yizhar O, Berndt A, Sohal VS, Deisseroth K, Hegemann P (March 2010). "Ultrafast optogenetic control".
241:
2527:
Zhang YP, Oertner TG (February 2007). "Optical induction of synaptic plasticity using a light-sensitive channel".
418:
Deisseroth's group has pioneered many applications in live animals such as genetically targeted remote control in
3221:
677:
114:
98:
retinal analogues, which led to the conclusion that the photoreceptor for motility responses in Chlorophyceae is
70:
5010:
5193:
4138:"NEUROSCIENCE. Natural light-gated anion channels: A family of microbial rhodopsins for advanced optogenetics"
2073:
Berndt A, Yizhar O, Gunaydin LA, Hegemann P, Deisseroth K (February 2009). "Bi-stable neural state switches".
1400:"NEUROSCIENCE. Natural light-gated anion channels: A family of microbial rhodopsins for advanced optogenetics"
1030:"The nature of rhodopsin-triggered photocurrents in Chlamydomonas. I. Kinetics and influence of divalent ions"
638:. Optical control of behavior has been demonstrated in nematodes, fruit flies, zebrafish, and mice. Recently,
5160:
3730:"An improved chloride-conducting channelrhodopsin for light-induced inhibition of neuronal activity in vivo"
245:
31:
4358:
Anisimova M, van Bommel B, Wang R, Mikhaylova M, Simon Wiegert J, Oertner TG, et al. (February 2022).
296:-retinal. This change introduces a further one in the transmembrane protein, opening the pore to at least 6
4412:
2670:"High-speed mapping of synaptic connectivity using photostimulation in Channelrhodopsin-2 transgenic mice"
2362:
701:
697:
428:
367:
229:
127:
3914:"Escape behavior elicited by single, channelrhodopsin-2-evoked spikes in zebrafish somatosensory neurons"
1457:"Kalium channelrhodopsins are natural light-gated potassium channels that mediate optogenetic inhibition"
622:
that behavior. Controlling networks of genetically modified cells with light, an emerging field known as
5183:
5142:
5033:
3847:
Fernandez Lahore RG, Pampaloni NP, Schiewer E, Heim MM, Tillert L, Vierock J, et al. (2022-12-21).
3520:
Fernandez Lahore RG, Pampaloni NP, Peter E, Heim MM, Tillert L, Vierock J, et al. (December 2022).
616:
this activity with behavior. This is not sufficient to prove that the recorded neural activity actually
437:
309:
4907:"In vivo light-induced activation of neural circuitry in transgenic mice expressing channelrhodopsin-2"
4301:"Optical induction of plasticity at single synapses reveals input-specific accumulation of alphaCaMKII"
684:
In 2006, it was reported that transfection with Channelrhodopsin could restore eyesight to blind mice.
3273:
Anisimova M, van Bommel B, Wang R, Mikhaylova M, Wiegert JS, Oertner TG, et al. (December 2022).
4778:
4471:
4312:
4255:
4149:
4094:
4039:
3982:
3925:
3860:
3801:
3741:
3681:
3621:
3533:
3076:
3017:
2958:
2856:
2797:
2740:
2681:
2478:
2421:
2354:
2297:
2240:
2183:
1980:
1793:
1731:
1620:
1525:
1411:
1316:
1208:
1098:
1041:
994:
943:
892:
736:
689:
200:
49:: movement in response to light. Expressed in cells of other organisms, they enable light to control
23:
4905:
Arenkiel BR, Peca J, Davison IG, Feliciano C, Deisseroth K, Augustine GJ, et al. (April 2007).
4822:
Keppeler D, Merino RM, Lopez de la Morena D, Bali B, Huet AT, Gehrt A, et al. (December 2018).
4765:
Mager T, Lopez de la Morena D, Senn V, Schlotte J, D Errico A, Feldbauer K, et al. (May 2018).
4417:
3124:"ReaChR: a red-shifted variant of channelrhodopsin enables deep transcranial optogenetic excitation"
3668:
Wietek J, Wiegert JS, Adeishvili N, Schneider F, Watanabe H, Tsunoda SP, et al. (April 2014).
3475:"Ultra light-sensitive and fast neuronal activation with the Ca²+-permeable channelrhodopsin CatCh"
2367:
673:
643:
478:
383:
321:
217:
1607:
Eria-Oliveira AS, Folacci M, Chassot AA, Fedou S, Thézé N, Zabelskii D, et al. (2024-01-02).
4656:
Sahel JA, Boulanger-Scemama E, Pagot C, Arleo A, Galluppi F, Martel JN, et al. (July 2021).
4638:
4440:
4118:
3971:"Sparse optical microstimulation in barrel cortex drives learned behaviour in freely moving mice"
3788:
Berndt A, Lee SY, Wietek J, Ramakrishnan C, Steinberg EE, Rashid AJ, et al. (January 2016).
3707:
3590:
3502:
2927:
2621:"The columnar and laminar organization of inhibitory connections to neocortical excitatory cells"
2598:
2552:
2098:
2052:
1895:
1700:
1589:
1494:
1010:
967:
916:
785:"Two rhodopsins mediate phototaxis to low- and high-intensity light in Chlamydomonas reinhardtii"
760:
589:
552:
4716:
Hernandez VH, Gehrt A, Reuter K, Jing Z, Jeschke M, Mendoza Schulz A, et al. (March 2014).
4242:
Vierock J, Rodriguez-Rozada S, Dieter A, Pieper F, Sims R, Tenedini F, et al. (July 2021).
3063:
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3004:
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2118:"Red-shifted optogenetic excitation: a tool for fast neural control derived from Volvox carteri"
1969:"Characterization of engineered channelrhodopsin variants with improved properties and kinetics"
372:
236:). The retinal chromophore is covalently linked to the rest of the protein through a protonated
4193:
Klapoetke NC, Murata Y, Kim SS, Pulver SR, Birdsey-Benson A, Cho YK, et al. (March 2014).
3171:
Klapoetke NC, Murata Y, Kim SS, Pulver SR, Birdsey-Benson A, Cho YK, et al. (March 2014).
2410:"Regulation of parkinsonian motor behaviours by optogenetic control of basal ganglia circuitry"
1303:
Kato HE, Zhang F, Yizhar O, Ramakrishnan C, Nishizawa T, Hirata K, et al. (January 2012).
1195:
Nagel G, Szellas T, Huhn W, Kateriya S, Adeishvili N, Berthold P, et al. (November 2003).
5090:
4980:
4936:
4892:
4853:
4804:
4747:
4679:
4630:
4595:
4546:
4497:
4430:
4389:
4340:
4281:
4224:
4175:
4110:
4067:
4008:
3951:
3894:
3876:
3829:
3767:
3699:
3647:
3559:
3494:
3455:
3404:
3353:
3304:
3255:
3202:
3153:
3104:
3045:
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2884:
2825:
2766:
2709:
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2504:
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2006:
1949:
1887:
1852:
1811:
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1654:
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1437:
1377:
1342:
1285:
1236:
1172:
1124:
1067:
959:
908:
865:
816:
752:
693:
400:
188:
75:
38:
5020:
4888:
3473:
Kleinlogel S, Feldbauer K, Dempski RE, Fotis H, Wood PG, Bamann C, et al. (April 2011).
5126:
4970:
4962:
4926:
4918:
4884:
4843:
4835:
4794:
4786:
4767:"High frequency neural spiking and auditory signaling by ultrafast red-shifted optogenetics"
4737:
4729:
4669:
4622:
4585:
4577:
4536:
4528:
4487:
4479:
4422:
4379:
4371:
4330:
4320:
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4263:
4214:
4206:
4165:
4157:
4102:
4057:
4047:
3998:
3990:
3969:
Huber D, Petreanu L, Ghitani N, Ranade S, Hromádka T, Mainen Z, et al. (January 2008).
3941:
3933:
3884:
3868:
3819:
3809:
3757:
3749:
3689:
3637:
3629:
3582:
3549:
3541:
3486:
3445:
3435:
3394:
3384:
3343:
3335:
3322:
Hochbaum DR, Zhao Y, Farhi SL, Klapoetke N, Werley CA, Kapoor V, et al. (August 2014).
3294:
3286:
3245:
3237:
3192:
3184:
3143:
3135:
3094:
3084:
3035:
3025:
2976:
2966:
2911:
2874:
2864:
2815:
2805:
2756:
2748:
2699:
2689:
2640:
2632:
2582:
2536:
2494:
2486:
2467:"Global and local fMRI signals driven by neurons defined optogenetically by type and wiring"
2437:
2429:
2380:
2372:
2313:
2305:
2256:
2248:
2199:
2191:
2172:"Neocortical excitation/inhibition balance in information processing and social dysfunction"
2170:
Yizhar O, Fenno LE, Prigge M, Schneider F, Davidson TJ, O'Shea DJ, et al. (July 2011).
2137:
2129:
2082:
2036:
1996:
1988:
1939:
1931:
1920:"A user's guide to channelrhodopsin variants: features, limitations and future developments"
1879:
1842:
1801:
1749:
1739:
1684:
1644:
1628:
1581:
1541:
1533:
1512:
Kim YS, Kato HE, Yamashita K, Ito S, Inoue K, Ramakrishnan C, et al. (September 2018).
1476:
1468:
1427:
1419:
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1332:
1324:
1275:
1267:
1226:
1216:
1164:
1114:
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1002:
951:
900:
855:
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806:
796:
744:
605:
482:
257:
50:
35:
3608:
Vierock J, Peter E, Grimm C, Rozenberg A, Chen IW, Tillert L, et al. (December 2022).
3226:"Dual-channel circuit mapping reveals sensorimotor convergence in the primary motor cortex"
5095:
5085:
4824:"Ultrafast optogenetic stimulation of the auditory pathway by targeting-optimized Chronos"
4244:"BiPOLES is an optogenetic tool developed for bidirectional dual-color control of neurons"
3790:"Structural foundations of optogenetics: Determinants of channelrhodopsin ion selectivity"
3324:"All-optical electrophysiology in mammalian neurons using engineered microbial rhodopsins"
2902:
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2408:
Kravitz AV, Freeze BS, Parker PR, Kay K, Thwin MT, Deisseroth K, et al. (July 2010).
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985:
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458:
4782:
4475:
4316:
4259:
4153:
4098:
4043:
3986:
3929:
3864:
3805:
3745:
3685:
3625:
3537:
3080:
3021:
2962:
2860:
2801:
2744:
2685:
2482:
2425:
2358:
2301:
2244:
2187:
2116:
Zhang F, Prigge M, Beyrière F, Tsunoda SP, Mattis J, Yizhar O, et al. (June 2008).
1984:
1797:
1735:
1649:
1624:
1608:
1529:
1415:
1320:
1212:
1102:
1045:
998:
947:
896:
740:
672:
can be identified. This is useful to study the molecular events during the induction of
653:
experiments. The blue-light sensitive ChR2 and the yellow light-activated chloride pump
4975:
4950:
4931:
4906:
4848:
4823:
4799:
4766:
4742:
4717:
4590:
4565:
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4516:
4492:
4459:
4384:
4359:
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4300:
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4243:
4219:
4194:
4170:
4137:
4062:
4027:
4003:
3970:
3946:
3913:
3889:
3848:
3824:
3789:
3762:
3729:
3642:
3609:
3554:
3521:
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3423:
3399:
3372:
3348:
3323:
3299:
3274:
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3225:
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3148:
3123:
3099:
3064:
3040:
3005:
2981:
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2844:
2820:
2785:
2761:
2728:
2704:
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2620:
2499:
2466:
2442:
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2385:
2342:
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2228:
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2171:
2142:
2117:
2001:
1968:
1944:
1919:
1780:
Nagel G, Brauner M, Liewald JF, Adeishvili N, Bamberg E, Gottschalk A (December 2005).
1754:
1719:
1546:
1513:
1481:
1456:
1432:
1399:
1337:
1304:
1280:
1255:
1119:
1086:
1062:
1029:
536:
387:
119:
3728:
Wietek J, Beltramo R, Scanziani M, Hegemann P, Oertner TG, Wiegert JS (October 2015).
2229:"Neural substrates of awakening probed with optogenetic control of hypocretin neurons"
1231:
1196:
1168:
1110:
1053:
860:
835:
811:
784:
477:
end of Channelrhodopsin-2 extends into the intracellular space and can be replaced by
5177:
5080:
3711:
3594:
3006:"High-efficiency channelrhodopsins for fast neuronal stimulation at low light levels"
2931:
1831:"Remote control of behavior through genetically targeted photostimulation of neurons"
1718:
Li X, Gutierrez DV, Hanson MG, Han J, Mark MD, Chiel H, et al. (December 2005).
1609:"Hijacking of internal calcium dynamics by intracellularly residing viral rhodopsins"
1593:
1498:
764:
654:
453:
for "the discovery of channelrhodopsin". In 2015, Boyden and Deisseroth received the
169:
106:
54:
5015:
5000:
4444:
3371:
Dana H, Mohar B, Sun Y, Narayan S, Gordus A, Hasseman JP, et al. (March 2016).
2668:
Wang H, Peca J, Matsuzaki M, Matsuzaki K, Noguchi J, Qiu L, et al. (May 2007).
2602:
2556:
2102:
1899:
1455:
Govorunova EG, Gou Y, Sineshchekov OA, Li H, Lu X, Wang Y, et al. (July 2022).
5056:
4642:
4122:
3506:
3241:
2845:"Two-photon single-cell optogenetic control of neuronal activity by sculpted light"
2727:
Mohanty SK, Reinscheid RK, Liu X, Okamura N, Krasieva TB, Berns MW (October 2008).
2465:
Lee JH, Durand R, Gradinaru V, Zhang F, Goshen I, Kim DS, et al. (June 2010).
2056:
1935:
1704:
1569:
Tajima S, Kim YS, Fukuda M, Byrne EF, Wang PY, Paggi JM, et al. (2022-10-31).
1014:
971:
920:
650:
631:
623:
577:
470:
325:
249:
68:). Channelrhodopsin-1 (ChR1) and Channelrhodopsin-2 (ChR2) from the model organism
65:
4658:"Partial recovery of visual function in a blind patient after optogenetic therapy"
427:, the optogenetic induction of learning in rodents, the experimental treatment of
4966:
4922:
4581:
4532:
4052:
2971:
2286:"Phasic firing in dopaminergic neurons is sufficient for behavioral conditioning"
851:
403:, permitting the control of individual action potentials at frequencies up to 200
3849:"Calcium-permeable channelrhodopsins for the photocontrol of calcium signalling"
3522:"Calcium-permeable channelrhodopsins for the photocontrol of calcium signalling"
3065:"Channelrhodopsin-2-XXL, a powerful optogenetic tool for low-light applications"
2752:
2227:
Adamantidis AR, Zhang F, Aravanis AM, Deisseroth K, de Lecea L (November 2007).
597:
517:
396:
237:
212:
42:
4949:
Bi A, Cui J, Ma YP, Olshevskaya E, Pu M, Dizhoor AM, et al. (April 2006).
4790:
4674:
4657:
4564:
Bi A, Cui J, Ma YP, Olshevskaya E, Pu M, Dizhoor AM, et al. (April 2006).
4515:
Bi A, Cui J, Ma YP, Olshevskaya E, Pu M, Dizhoor AM, et al. (April 2006).
4305:
Proceedings of the National Academy of Sciences of the United States of America
4267:
3872:
3794:
Proceedings of the National Academy of Sciences of the United States of America
3545:
3069:
Proceedings of the National Academy of Sciences of the United States of America
3010:
Proceedings of the National Academy of Sciences of the United States of America
2915:
2849:
Proceedings of the National Academy of Sciences of the United States of America
2790:
Proceedings of the National Academy of Sciences of the United States of America
2674:
Proceedings of the National Academy of Sciences of the United States of America
1847:
1830:
1724:
Proceedings of the National Academy of Sciences of the United States of America
1632:
1571:"Structural basis for ion selectivity in potassium-selective channelrhodopsins"
1472:
1254:
Berthold P, Tsunoda SP, Ernst OP, Mages W, Gradmann D, Hegemann P (June 2008).
1201:
Proceedings of the National Academy of Sciences of the United States of America
789:
Proceedings of the National Academy of Sciences of the United States of America
3937:
3586:
1992:
1806:
1781:
1585:
1537:
1373:
1197:"Channelrhodopsin-2, a directly light-gated cation-selective membrane channel"
1087:"Two light-activated conductances in the eye of the green alga Volvox carteri"
688:
delivered to the eyes of a human patient suffering from retinal degeneration (
474:
355:
253:
94:
46:
4426:
4375:
3880:
3440:
3290:
1640:
4839:
4325:
4161:
3814:
3694:
3669:
3089:
3030:
2869:
2810:
2694:
2619:
Kätzel D, Zemelman BV, Buetfering C, Wölfel M, Miesenböck G (January 2011).
2376:
2341:
Gradinaru V, Mogri M, Thompson KR, Henderson JM, Deisseroth K (April 2009).
2309:
1744:
1423:
1221:
748:
627:
618:
601:
359:
233:
208:
192:
99:
27:
4984:
4940:
4896:
4857:
4808:
4751:
4698:
4683:
4634:
4599:
4550:
4501:
4393:
4360:"Spike-timing-dependent plasticity rewards synchrony rather than causality"
4344:
4285:
4228:
4179:
4114:
4071:
4012:
3955:
3898:
3833:
3771:
3703:
3651:
3633:
3563:
3498:
3459:
3422:
Afshar Saber W, Gasparoli FM, Dirks MG, Gunn-Moore FJ, Antkowiak M (2018).
3408:
3357:
3308:
3275:"Spike-timing-dependent plasticity rewards synchrony rather than causality"
3259:
3206:
3157:
3108:
3049:
2990:
2923:
2888:
2829:
2770:
2713:
2654:
2594:
2548:
2508:
2451:
2394:
2327:
2270:
2213:
2151:
2094:
2048:
2010:
1953:
1891:
1856:
1815:
1763:
1696:
1658:
1555:
1514:"Crystal structure of the natural anion-conducting channelrhodopsin GtACR1"
1490:
1441:
1381:
1346:
1289:
1271:
1240:
1176:
1128:
820:
801:
756:
547:
the membrane upon illumination, preventing spike generation (inhibitory).
138:
by three research groups generated confusion about their naming: The names
1071:
1028:
Holland EM, Braun FJ, Nonnengässer C, Harz H, Hegemann P (February 1996).
912:
869:
4901:(Naturel function of channelrhodopsins and other photoreceptors in green)
4136:
Govorunova EG, Sineshchekov OA, Janz R, Liu X, Spudich JL (August 2015).
1398:
Govorunova EG, Sineshchekov OA, Janz R, Liu X, Spudich JL (August 2015).
963:
705:
669:
585:
345:
276:
The natural ("wild-type") ChR2 absorbs blue light with an absorption and
226:
4483:
4106:
3994:
3912:
Douglass AD, Kraves S, Deisseroth K, Schier AF, Engert F (August 2008).
3389:
2490:
2433:
2252:
2195:
1328:
4210:
3339:
3188:
593:
573:
540:
423:
419:
350:
268:
222:
164:
135:
83:
61:
57:
3753:
3373:"Sensitive red protein calcium indicators for imaging neural activity"
1305:"Crystal structure of the channelrhodopsin light-gated cation channel"
572:
Channelrhodopsins can be readily expressed in excitable cells such as
4733:
2540:
1883:
1006:
955:
904:
340:
329:
301:
285:
4945:(Using channelrhodopsin in transgenic mice to study brain circuitry)
4626:
4460:"The subcellular organization of neocortical excitatory connections"
3670:"Conversion of channelrhodopsin into a light-gated chloride channel"
3490:
3139:
3122:
Lin JY, Knutsen PM, Muller A, Kleinfeld D, Tsien RY (October 2013).
2636:
2133:
2086:
2040:
118:
were studied over many years in the groups of Oleg Sineshchekov and
3474:
2586:
1688:
543:. When expressed in neurons, potassium-selective channelrhodopsins
445:
excitation of ChR2, permitting the activation of individual cells.
665:
320:
In 2005, three groups sequentially established ChR2 as a tool for
267:
182:
79:
523:
A blue-shifted channelrhodopsin has been discovered in the alga
432:
252:), channelrhodopsins directly form ion channels (i.e., they are
5029:
5025:
4195:"Independent optical excitation of distinct neural populations"
3173:"Independent optical excitation of distinct neural populations"
183:
2947:"Temporal control of immediate early gene induction by light"
649:
Using multiple colors of light expands the possibilities of
4458:
Petreanu L, Mao T, Sternson SM, Svoboda K (February 2009).
2843:
Andrasfalvy BK, Zemelman BV, Tang J, Vaziri A (June 2010).
2786:"Two-photon excitation of channelrhodopsin-2 at saturation"
339:'s lab demonstrated that ChR2 could be deployed to control
5005:
2343:"Optical deconstruction of parkinsonian neural circuitry"
630:
link between activity in a specific group of neurons and
516:
minimally invasive, transcranial activation of brainstem
457:
and in 2020, Miesenböck, Hegemann and Nagel received the
2945:
Schoenenberger P, Gerosa D, Oertner TG (December 2009).
4989:(Using channelrhodopsin potentially to treat blindness)
3424:"All-Optical Assay to Study Biological Neural Networks"
692:), leading to partial recovery of his vision. Optical
146:
were used for initial submission by Hegemann's group;
2522:
2520:
2518:
1967:
Lin JY, Lin MZ, Steinbach P, Tsien RY (March 2009).
304:channels (CCRs), conducting H, Na, K, and Ca ions.
5135:
5104:
5063:
4875:Hegemann P (2008). "Algal sensory photoreceptors".
1157:
Biochemical and Biophysical Research Communications
778:
776:
774:
664:Using fluorescently labeled ChR2, light-stimulated
1393:
1391:
783:Sineshchekov OA, Jung KH, Spudich JL (June 2002).
576:using a variety of transfection techniques (viral
134:. Independent submission of the same sequences to
4718:"Optogenetic stimulation of the auditory pathway"
4411:. IEEE Decision and Control. pp. 6798–6805.
481:without affecting channel function. This kind of
2568:
2566:
2022:
2020:
312:and lead to calcium release when illuminated.
4699:"Algae proteins partially restore man's sight"
4299:Zhang YP, Holbro N, Oertner TG (August 2008).
3783:
3781:
3723:
3721:
3663:
3661:
2614:
2612:
2165:
2163:
2161:
2068:
2066:
1670:
1668:
288:, it induces a conformational change from all-
5041:
5001:Optogenetics Resource Center / Deisseroth lab
1775:
1773:
1190:
1188:
1186:
199:In terms of structure, channelrhodopsins are
8:
4409:52nd IEEE Conference on Decision and Control
722:
720:
244:that open other ion channels indirectly via
240:. Whereas most 7-transmembrane proteins are
1913:
1911:
1909:
5048:
5034:
5026:
64:influx, and other cellular processes (see
4974:
4930:
4847:
4798:
4741:
4673:
4589:
4540:
4491:
4416:
4383:
4334:
4324:
4275:
4218:
4169:
4061:
4051:
4002:
3945:
3888:
3823:
3813:
3761:
3693:
3641:
3553:
3449:
3439:
3398:
3388:
3347:
3298:
3249:
3196:
3147:
3098:
3088:
3039:
3029:
2980:
2970:
2878:
2868:
2819:
2809:
2760:
2703:
2693:
2644:
2498:
2441:
2384:
2366:
2317:
2260:
2203:
2141:
2000:
1943:
1846:
1805:
1753:
1743:
1648:
1545:
1480:
1431:
1336:
1279:
1230:
1220:
1118:
1061:
859:
810:
800:
626:., allows researchers now to explore the
612:the activity of neurons in the brain and
4889:10.1146/annurev.arplant.59.032607.092847
2784:Rickgauer JP, Tank DW (September 2009).
704:patients or to control certain forms of
716:
459:Shaw prize in Life Science and Medicine
187:Crystal structure of channelrhodopsin.
441:conditional paradigm – as well as the
4722:The Journal of Clinical Investigation
836:"Light Antennas in phototactic algae"
834:Foster KW, Smyth RD (December 1980).
640:chloride-conducting channelrhodopsins
461:for the development of optogenetics.
280:maximum at 480 nm. When the all-
211:, and contain the light-isomerizable
7:
5156:
1829:Lima SQ, Miesenböck G (April 2005).
1085:Braun FJ, Hegemann P (March 1999).
469:Channelrhodopsins are key tools in
455:Breakthrough Prize in Life Sciences
272:Scheme of ChR2-RFP fusion construct
557:Anion-conducting channelrhodopsins
431:in rats, and the combination with
306:Anion-conducting channelrhodopsins
14:
324:targeted optical remote control (
5155:
451:Louis-Jeantet Prize for Medicine
407:Hz (in appropriate cell types).
332:, neural circuits and behavior.
4026:Han X, Boyden ES (March 2007).
316:Development as a molecular tool
4877:Annual Review of Plant Biology
3242:10.1523/JNEUROSCI.3741-14.2015
1936:10.1113/expphysiol.2009.051961
592:. The light-absorbing pigment
1:
1169:10.1016/S0006-291X(02)03079-6
1111:10.1016/S0006-3495(99)77326-1
1054:10.1016/S0006-3495(96)79635-2
596:is present in most cells (of
4967:10.1016/j.neuron.2006.02.026
4923:10.1016/j.neuron.2007.03.005
4582:10.1016/j.neuron.2006.02.026
4533:10.1016/j.neuron.2006.02.026
4053:10.1371/journal.pone.0000299
2972:10.1371/journal.pone.0008185
1362:Journal of Molecular Biology
852:10.1128/mr.44.4.572-630.1980
205:seven-transmembrane proteins
4697:Gallagher J (24 May 2021).
3230:The Journal of Neuroscience
2753:10.1529/biophysj.108.130187
284:-retinal complex absorbs a
242:G protein-coupled receptors
16:Class of transport proteins
5210:
5189:Integral membrane proteins
4791:10.1038/s41467-018-04146-3
4675:10.1038/s41591-021-01351-4
4268:10.1038/s41467-021-24759-5
3873:10.1038/s41467-022-35373-4
3546:10.1038/s41467-022-35373-4
2916:10.1016/j.tins.2013.08.005
1848:10.1016/j.cell.2005.02.004
1633:10.1038/s41467-023-44548-6
1473:10.1038/s41593-022-01094-6
678:cultured neuronal networks
465:Designer-channelrhodopsins
5151:
3938:10.1016/j.cub.2008.06.077
3587:10.1101/2021.09.17.460684
3428:Frontiers in Neuroscience
3220:Hooks BM, Lin JY, Guo C,
1993:10.1016/j.bpj.2008.11.034
1807:10.1016/j.cub.2005.11.032
1586:10.1101/2022.10.30.514430
1538:10.1038/s41586-018-0511-6
1374:10.1016/j.jmb.2007.10.072
115:Chlamydomonas reinhardtii
71:Chlamydomonas reinhardtii
4427:10.1109/CDC.2013.6760966
3441:10.3389/fnins.2018.00451
604:, making it possible to
154:by Spudich's group; and
32:light-gated ion channels
4840:10.15252/embj.201899649
4326:10.1073/pnas.0802940105
4162:10.1126/science.aaa7484
3815:10.1073/pnas.1523341113
3695:10.1126/science.1249375
3090:10.1073/pnas.1408269111
3031:10.1073/pnas.1017210108
2904:Trends in Neurosciences
2870:10.1073/pnas.1006620107
2811:10.1073/pnas.0907084106
2695:10.1073/pnas.0700384104
2377:10.1126/science.1167093
2310:10.1126/science.1168878
1924:Experimental Physiology
1918:Lin JY (January 2011).
1745:10.1073/pnas.0509030102
1424:10.1126/science.aaa7484
1222:10.1073/pnas.1936192100
840:Microbiological Reviews
749:10.1126/science.1072068
110:Haematococcus pluvialis
51:electrical excitability
4376:10.1093/cercor/bhac050
3634:10.1126/sciadv.add7729
3291:10.1093/cercor/bhac050
1272:10.1105/tpc.108.057919
802:10.1073/pnas.122243399
698:deep-brain stimulation
502:Photocurrent amplitude
273:
196:
130:sequencing project in
78:from other species of
5143:Voltage-sensitive dye
5064:Optogenetic actuators
4771:Nature Communications
4248:Nature Communications
3853:Nature Communications
3581:: 2021.09.17.460684.
3526:Nature Communications
1613:Nature Communications
310:endoplasmic reticulum
271:
201:retinylidene proteins
186:
105:Photocurrents of the
24:retinylidene proteins
690:retinitis pigmentosa
479:fluorescent proteins
5105:Optogenetic sensors
4783:2018NatCo...9.1750M
4615:Nature Neuroscience
4484:10.1038/nature07709
4476:2009Natur.457.1142P
4470:(7233): 1142–1145.
4317:2008PNAS..10512039Z
4311:(33): 12039–12044.
4260:2021NatCo..12.4527V
4154:2015Sci...349..647G
4107:10.1038/nature05744
4099:2007Natur.446..633Z
4044:2007PLoSO...2..299H
3995:10.1038/nature06445
3987:2008Natur.451...61H
3930:2008CBio...18.1133D
3865:2022NatCo..13.7844F
3806:2016PNAS..113..822B
3746:2015NatSR...514807W
3686:2014Sci...344..409W
3626:2022SciA....8D7729V
3538:2022NatCo..13.7844F
3479:Nature Neuroscience
3390:10.7554/eLife.12727
3128:Nature Neuroscience
3081:2014PNAS..11113972D
3075:(38): 13972–13977.
3022:2011PNAS..108.7595B
2963:2009PLoSO...4.8185S
2861:2010PNAS..10711981A
2855:(26): 11981–11986.
2802:2009PNAS..10615025R
2796:(35): 15025–15030.
2745:2008BpJ....95.3916M
2733:Biophysical Journal
2686:2007PNAS..104.8143W
2625:Nature Neuroscience
2575:Nature Neuroscience
2491:10.1038/nature09108
2483:2010Natur.465..788L
2434:10.1038/nature09159
2426:2010Natur.466..622K
2359:2009Sci...324..354G
2302:2009Sci...324.1080T
2296:(5930): 1080–1084.
2253:10.1038/nature06310
2245:2007Natur.450..420A
2196:10.1038/nature10360
2188:2011Natur.477..171Y
2122:Nature Neuroscience
2075:Nature Neuroscience
2029:Nature Neuroscience
1985:2009BpJ....96.1803L
1973:Biophysical Journal
1798:2005CBio...15.2279N
1736:2005PNAS..10217816L
1730:(49): 17816–17821.
1677:Nature Neuroscience
1625:2024NatCo..15...65E
1530:2018Natur.561..343K
1461:Nature Neuroscience
1416:2015Sci...349..647G
1329:10.1038/nature10870
1321:2012Natur.482..369K
1213:2003PNAS..10013940N
1207:(24): 13940–13945.
1103:1999BpJ....76.1668B
1091:Biophysical Journal
1046:1996BpJ....70..924H
1034:Biophysical Journal
999:1991Natur.351..489H
948:1978Natur.271..476L
897:1984Natur.311..756F
741:2002Sci...296.2395N
735:(5577): 2395–2398.
674:synaptic plasticity
644:shunting inhibition
429:Parkinson's disease
384:genetic engineering
132:C. reinhardtii
30:) that function as
22:are a subfamily of
5136:Related techniques
4211:10.1038/nmeth.2836
3734:Scientific Reports
3340:10.1038/nmeth.3000
3189:10.1038/nmeth.2836
590:transgenic animals
553:reversal potential
274:
197:
170:human kidney cells
5171:
5170:
5091:Bacteriorhodopsin
4436:978-1-4673-5717-3
4148:(6248): 647–650.
4093:(7136): 633–639.
3924:(15): 1133–1137.
3754:10.1038/srep14807
3680:(6182): 409–412.
3236:(10): 4418–4426.
3134:(10): 1499–1508.
3016:(18): 7595–7600.
2680:(19): 8143–8148.
2477:(7299): 788–792.
2420:(7306): 622–626.
2353:(5925): 354–359.
2239:(7168): 420–424.
2182:(7363): 171–178.
1792:(24): 2279–2284.
1524:(7723): 343–348.
1410:(6248): 647–650.
1315:(7385): 369–374.
993:(6326): 489–491.
942:(5644): 476–478.
891:(5988): 756–759.
795:(13): 8689–8694.
694:cochlear implants
525:Scherffelia dubia
246:second messengers
20:Channelrhodopsins
5201:
5159:
5158:
5075:Anion-conducting
5071:Channelrhodopsin
5050:
5043:
5036:
5027:
4988:
4978:
4944:
4934:
4900:
4862:
4861:
4851:
4828:The EMBO Journal
4819:
4813:
4812:
4802:
4762:
4756:
4755:
4745:
4734:10.1172/JCI69050
4728:(3): 1114–1129.
4713:
4707:
4706:
4694:
4688:
4687:
4677:
4668:(7): 1223–1229.
4653:
4647:
4646:
4610:
4604:
4603:
4593:
4561:
4555:
4554:
4544:
4512:
4506:
4505:
4495:
4455:
4449:
4448:
4420:
4404:
4398:
4397:
4387:
4355:
4349:
4348:
4338:
4328:
4296:
4290:
4289:
4279:
4239:
4233:
4232:
4222:
4190:
4184:
4183:
4173:
4133:
4127:
4126:
4082:
4076:
4075:
4065:
4055:
4023:
4017:
4016:
4006:
3966:
3960:
3959:
3949:
3909:
3903:
3902:
3892:
3844:
3838:
3837:
3827:
3817:
3785:
3776:
3775:
3765:
3725:
3716:
3715:
3697:
3665:
3656:
3655:
3645:
3620:(49): eadd7729.
3614:Science Advances
3605:
3599:
3598:
3574:
3568:
3567:
3557:
3517:
3511:
3510:
3470:
3464:
3463:
3453:
3443:
3419:
3413:
3412:
3402:
3392:
3368:
3362:
3361:
3351:
3319:
3313:
3312:
3302:
3270:
3264:
3263:
3253:
3217:
3211:
3210:
3200:
3168:
3162:
3161:
3151:
3119:
3113:
3112:
3102:
3092:
3060:
3054:
3053:
3043:
3033:
3001:
2995:
2994:
2984:
2974:
2942:
2936:
2935:
2899:
2893:
2892:
2882:
2872:
2840:
2834:
2833:
2823:
2813:
2781:
2775:
2774:
2764:
2739:(8): 3916–3926.
2724:
2718:
2717:
2707:
2697:
2665:
2659:
2658:
2648:
2616:
2607:
2606:
2570:
2561:
2560:
2541:10.1038/nmeth988
2524:
2513:
2512:
2502:
2462:
2456:
2455:
2445:
2405:
2399:
2398:
2388:
2370:
2338:
2332:
2331:
2321:
2281:
2275:
2274:
2264:
2224:
2218:
2217:
2207:
2167:
2156:
2155:
2145:
2113:
2107:
2106:
2070:
2061:
2060:
2024:
2015:
2014:
2004:
1979:(5): 1803–1814.
1964:
1958:
1957:
1947:
1915:
1904:
1903:
1884:10.1038/nmeth936
1867:
1861:
1860:
1850:
1826:
1820:
1819:
1809:
1777:
1768:
1767:
1757:
1747:
1715:
1709:
1708:
1683:(9): 1263–1268.
1672:
1663:
1662:
1652:
1604:
1598:
1597:
1575:
1566:
1560:
1559:
1549:
1509:
1503:
1502:
1484:
1452:
1446:
1445:
1435:
1395:
1386:
1385:
1357:
1351:
1350:
1340:
1300:
1294:
1293:
1283:
1266:(6): 1665–1677.
1251:
1245:
1244:
1234:
1224:
1192:
1181:
1180:
1152:
1146:
1143:accession number
1139:
1133:
1132:
1122:
1097:(3): 1668–1678.
1082:
1076:
1075:
1065:
1025:
1019:
1018:
1007:10.1038/351489a0
982:
976:
975:
956:10.1038/271476a0
931:
925:
924:
905:10.1038/311756a0
880:
874:
873:
863:
831:
825:
824:
814:
804:
780:
769:
768:
724:
555:more negative.
483:fusion construct
258:photostimulation
248:(i.e., they are
195:
34:. They serve as
5209:
5208:
5204:
5203:
5202:
5200:
5199:
5198:
5194:Neurotechnology
5174:
5173:
5172:
5167:
5147:
5131:
5100:
5096:Proteorhodopsin
5086:Archaerhodopsin
5059:
5054:
4997:
4992:
4948:
4904:
4874:
4870:
4868:Further reading
4865:
4821:
4820:
4816:
4764:
4763:
4759:
4715:
4714:
4710:
4696:
4695:
4691:
4662:Nature Medicine
4655:
4654:
4650:
4627:10.1038/nn.2117
4612:
4611:
4607:
4563:
4562:
4558:
4514:
4513:
4509:
4457:
4456:
4452:
4437:
4418:10.1.1.671.6351
4406:
4405:
4401:
4364:Cerebral Cortex
4357:
4356:
4352:
4298:
4297:
4293:
4241:
4240:
4236:
4192:
4191:
4187:
4135:
4134:
4130:
4084:
4083:
4079:
4025:
4024:
4020:
3981:(7174): 61–64.
3968:
3967:
3963:
3918:Current Biology
3911:
3910:
3906:
3846:
3845:
3841:
3787:
3786:
3779:
3727:
3726:
3719:
3667:
3666:
3659:
3607:
3606:
3602:
3576:
3575:
3571:
3519:
3518:
3514:
3491:10.1038/nn.2776
3472:
3471:
3467:
3421:
3420:
3416:
3370:
3369:
3365:
3321:
3320:
3316:
3279:Cerebral Cortex
3272:
3271:
3267:
3219:
3218:
3214:
3170:
3169:
3165:
3140:10.1038/nn.3502
3121:
3120:
3116:
3062:
3061:
3057:
3003:
3002:
2998:
2944:
2943:
2939:
2910:(10): 557–560.
2901:
2900:
2896:
2842:
2841:
2837:
2783:
2782:
2778:
2726:
2725:
2721:
2667:
2666:
2662:
2637:10.1038/nn.2687
2618:
2617:
2610:
2572:
2571:
2564:
2526:
2525:
2516:
2464:
2463:
2459:
2407:
2406:
2402:
2340:
2339:
2335:
2283:
2282:
2278:
2226:
2225:
2221:
2169:
2168:
2159:
2134:10.1038/nn.2120
2115:
2114:
2110:
2087:10.1038/nn.2247
2072:
2071:
2064:
2041:10.1038/nn.2495
2026:
2025:
2018:
1966:
1965:
1961:
1917:
1916:
1907:
1878:(10): 785–792.
1869:
1868:
1864:
1828:
1827:
1823:
1786:Current Biology
1779:
1778:
1771:
1717:
1716:
1712:
1674:
1673:
1666:
1606:
1605:
1601:
1573:
1568:
1567:
1563:
1511:
1510:
1506:
1454:
1453:
1449:
1397:
1396:
1389:
1359:
1358:
1354:
1302:
1301:
1297:
1253:
1252:
1248:
1194:
1193:
1184:
1154:
1153:
1149:
1140:
1136:
1084:
1083:
1079:
1027:
1026:
1022:
984:
983:
979:
933:
932:
928:
882:
881:
877:
833:
832:
828:
782:
781:
772:
726:
725:
718:
714:
676:. Transfected
636:decision making
582:electroporation
570:
559:(iChloC, iC++,
533:
531:Ion selectivity
513:
504:
495:
487:Point mutations
467:
393:C. elegans
373:Gero Miesenböck
368:C. elegans
337:Karl Deisseroth
318:
278:action spectrum
266:
191:
181:
92:
41:in unicellular
17:
12:
11:
5:
5207:
5205:
5197:
5196:
5191:
5186:
5176:
5175:
5169:
5168:
5166:
5165:
5152:
5149:
5148:
5146:
5145:
5139:
5137:
5133:
5132:
5130:
5129:
5124:
5119:
5114:
5108:
5106:
5102:
5101:
5099:
5098:
5093:
5088:
5083:
5078:
5067:
5065:
5061:
5060:
5055:
5053:
5052:
5045:
5038:
5030:
5024:
5023:
5018:
5013:
5008:
5003:
4996:
4995:External links
4993:
4991:
4990:
4946:
4917:(2): 205–218.
4902:
4871:
4869:
4866:
4864:
4863:
4834:(24): e99649.
4814:
4757:
4708:
4689:
4648:
4621:(6): 667–675.
4605:
4556:
4507:
4450:
4435:
4399:
4350:
4291:
4234:
4205:(3): 338–346.
4199:Nature Methods
4185:
4128:
4077:
4018:
3961:
3904:
3839:
3800:(4): 822–829.
3777:
3717:
3657:
3600:
3569:
3512:
3485:(4): 513–518.
3465:
3414:
3363:
3334:(8): 825–833.
3328:Nature Methods
3314:
3265:
3224:(March 2015).
3212:
3183:(3): 338–346.
3177:Nature Methods
3163:
3114:
3055:
2996:
2937:
2894:
2835:
2776:
2719:
2660:
2631:(1): 100–107.
2608:
2587:10.1038/nn1891
2581:(5): 663–668.
2562:
2535:(2): 139–141.
2529:Nature Methods
2514:
2457:
2400:
2368:10.1.1.368.668
2333:
2276:
2219:
2157:
2128:(6): 631–633.
2108:
2081:(2): 229–234.
2062:
2035:(3): 387–392.
2016:
1959:
1905:
1872:Nature Methods
1862:
1841:(1): 141–152.
1821:
1769:
1710:
1689:10.1038/nn1525
1664:
1599:
1561:
1504:
1467:(7): 967–974.
1447:
1387:
1368:(3): 686–694.
1352:
1295:
1260:The Plant Cell
1246:
1182:
1163:(3): 711–717.
1147:
1134:
1077:
1040:(2): 924–931.
1020:
977:
926:
875:
846:(4): 572–630.
826:
770:
715:
713:
710:
606:photostimulate
569:
566:
532:
529:
512:
509:
503:
500:
494:
491:
466:
463:
413:Volvox carteri
388:point mutation
317:
314:
265:
262:
180:
177:
120:Peter Hegemann
91:
88:
45:, controlling
39:photoreceptors
15:
13:
10:
9:
6:
4:
3:
2:
5206:
5195:
5192:
5190:
5187:
5185:
5182:
5181:
5179:
5164:
5163:
5154:
5153:
5150:
5144:
5141:
5140:
5138:
5134:
5128:
5125:
5123:
5120:
5118:
5115:
5113:
5110:
5109:
5107:
5103:
5097:
5094:
5092:
5089:
5087:
5084:
5082:
5081:Halorhodopsin
5079:
5076:
5072:
5069:
5068:
5066:
5062:
5058:
5051:
5046:
5044:
5039:
5037:
5032:
5031:
5028:
5022:
5019:
5017:
5014:
5012:
5009:
5007:
5004:
5002:
4999:
4998:
4994:
4986:
4982:
4977:
4972:
4968:
4964:
4960:
4956:
4952:
4947:
4942:
4938:
4933:
4928:
4924:
4920:
4916:
4912:
4908:
4903:
4898:
4894:
4890:
4886:
4882:
4878:
4873:
4872:
4867:
4859:
4855:
4850:
4845:
4841:
4837:
4833:
4829:
4825:
4818:
4815:
4810:
4806:
4801:
4796:
4792:
4788:
4784:
4780:
4776:
4772:
4768:
4761:
4758:
4753:
4749:
4744:
4739:
4735:
4731:
4727:
4723:
4719:
4712:
4709:
4704:
4700:
4693:
4690:
4685:
4681:
4676:
4671:
4667:
4663:
4659:
4652:
4649:
4644:
4640:
4636:
4632:
4628:
4624:
4620:
4616:
4609:
4606:
4601:
4597:
4592:
4587:
4583:
4579:
4575:
4571:
4567:
4560:
4557:
4552:
4548:
4543:
4538:
4534:
4530:
4526:
4522:
4518:
4511:
4508:
4503:
4499:
4494:
4489:
4485:
4481:
4477:
4473:
4469:
4465:
4461:
4454:
4451:
4446:
4442:
4438:
4432:
4428:
4424:
4419:
4414:
4410:
4403:
4400:
4395:
4391:
4386:
4381:
4377:
4373:
4369:
4365:
4361:
4354:
4351:
4346:
4342:
4337:
4332:
4327:
4322:
4318:
4314:
4310:
4306:
4302:
4295:
4292:
4287:
4283:
4278:
4273:
4269:
4265:
4261:
4257:
4253:
4249:
4245:
4238:
4235:
4230:
4226:
4221:
4216:
4212:
4208:
4204:
4200:
4196:
4189:
4186:
4181:
4177:
4172:
4167:
4163:
4159:
4155:
4151:
4147:
4143:
4139:
4132:
4129:
4124:
4120:
4116:
4112:
4108:
4104:
4100:
4096:
4092:
4088:
4081:
4078:
4073:
4069:
4064:
4059:
4054:
4049:
4045:
4041:
4037:
4033:
4029:
4022:
4019:
4014:
4010:
4005:
4000:
3996:
3992:
3988:
3984:
3980:
3976:
3972:
3965:
3962:
3957:
3953:
3948:
3943:
3939:
3935:
3931:
3927:
3923:
3919:
3915:
3908:
3905:
3900:
3896:
3891:
3886:
3882:
3878:
3874:
3870:
3866:
3862:
3858:
3854:
3850:
3843:
3840:
3835:
3831:
3826:
3821:
3816:
3811:
3807:
3803:
3799:
3795:
3791:
3784:
3782:
3778:
3773:
3769:
3764:
3759:
3755:
3751:
3747:
3743:
3739:
3735:
3731:
3724:
3722:
3718:
3713:
3709:
3705:
3701:
3696:
3691:
3687:
3683:
3679:
3675:
3671:
3664:
3662:
3658:
3653:
3649:
3644:
3639:
3635:
3631:
3627:
3623:
3619:
3615:
3611:
3604:
3601:
3596:
3592:
3588:
3584:
3580:
3573:
3570:
3565:
3561:
3556:
3551:
3547:
3543:
3539:
3535:
3531:
3527:
3523:
3516:
3513:
3508:
3504:
3500:
3496:
3492:
3488:
3484:
3480:
3476:
3469:
3466:
3461:
3457:
3452:
3447:
3442:
3437:
3433:
3429:
3425:
3418:
3415:
3410:
3406:
3401:
3396:
3391:
3386:
3382:
3378:
3374:
3367:
3364:
3359:
3355:
3350:
3345:
3341:
3337:
3333:
3329:
3325:
3318:
3315:
3310:
3306:
3301:
3296:
3292:
3288:
3284:
3280:
3276:
3269:
3266:
3261:
3257:
3252:
3247:
3243:
3239:
3235:
3231:
3227:
3223:
3216:
3213:
3208:
3204:
3199:
3194:
3190:
3186:
3182:
3178:
3174:
3167:
3164:
3159:
3155:
3150:
3145:
3141:
3137:
3133:
3129:
3125:
3118:
3115:
3110:
3106:
3101:
3096:
3091:
3086:
3082:
3078:
3074:
3070:
3066:
3059:
3056:
3051:
3047:
3042:
3037:
3032:
3027:
3023:
3019:
3015:
3011:
3007:
3000:
2997:
2992:
2988:
2983:
2978:
2973:
2968:
2964:
2960:
2957:(12): e8185.
2956:
2952:
2948:
2941:
2938:
2933:
2929:
2925:
2921:
2917:
2913:
2909:
2905:
2898:
2895:
2890:
2886:
2881:
2876:
2871:
2866:
2862:
2858:
2854:
2850:
2846:
2839:
2836:
2831:
2827:
2822:
2817:
2812:
2807:
2803:
2799:
2795:
2791:
2787:
2780:
2777:
2772:
2768:
2763:
2758:
2754:
2750:
2746:
2742:
2738:
2734:
2730:
2723:
2720:
2715:
2711:
2706:
2701:
2696:
2691:
2687:
2683:
2679:
2675:
2671:
2664:
2661:
2656:
2652:
2647:
2642:
2638:
2634:
2630:
2626:
2622:
2615:
2613:
2609:
2604:
2600:
2596:
2592:
2588:
2584:
2580:
2576:
2569:
2567:
2563:
2558:
2554:
2550:
2546:
2542:
2538:
2534:
2530:
2523:
2521:
2519:
2515:
2510:
2506:
2501:
2496:
2492:
2488:
2484:
2480:
2476:
2472:
2468:
2461:
2458:
2453:
2449:
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2427:
2423:
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2415:
2411:
2404:
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2374:
2369:
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2360:
2356:
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2311:
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2299:
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2263:
2258:
2254:
2250:
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2238:
2234:
2230:
2223:
2220:
2215:
2211:
2206:
2201:
2197:
2193:
2189:
2185:
2181:
2177:
2173:
2166:
2164:
2162:
2158:
2153:
2149:
2144:
2139:
2135:
2131:
2127:
2123:
2119:
2112:
2109:
2104:
2100:
2096:
2092:
2088:
2084:
2080:
2076:
2069:
2067:
2063:
2058:
2054:
2050:
2046:
2042:
2038:
2034:
2030:
2023:
2021:
2017:
2012:
2008:
2003:
1998:
1994:
1990:
1986:
1982:
1978:
1974:
1970:
1963:
1960:
1955:
1951:
1946:
1941:
1937:
1933:
1929:
1925:
1921:
1914:
1912:
1910:
1906:
1901:
1897:
1893:
1889:
1885:
1881:
1877:
1873:
1866:
1863:
1858:
1854:
1849:
1844:
1840:
1836:
1832:
1825:
1822:
1817:
1813:
1808:
1803:
1799:
1795:
1791:
1787:
1783:
1776:
1774:
1770:
1765:
1761:
1756:
1751:
1746:
1741:
1737:
1733:
1729:
1725:
1721:
1714:
1711:
1706:
1702:
1698:
1694:
1690:
1686:
1682:
1678:
1671:
1669:
1665:
1660:
1656:
1651:
1646:
1642:
1638:
1634:
1630:
1626:
1622:
1618:
1614:
1610:
1603:
1600:
1595:
1591:
1587:
1583:
1579:
1572:
1565:
1562:
1557:
1553:
1548:
1543:
1539:
1535:
1531:
1527:
1523:
1519:
1515:
1508:
1505:
1500:
1496:
1492:
1488:
1483:
1478:
1474:
1470:
1466:
1462:
1458:
1451:
1448:
1443:
1439:
1434:
1429:
1425:
1421:
1417:
1413:
1409:
1405:
1401:
1394:
1392:
1388:
1383:
1379:
1375:
1371:
1367:
1363:
1356:
1353:
1348:
1344:
1339:
1334:
1330:
1326:
1322:
1318:
1314:
1310:
1306:
1299:
1296:
1291:
1287:
1282:
1277:
1273:
1269:
1265:
1261:
1257:
1250:
1247:
1242:
1238:
1233:
1228:
1223:
1218:
1214:
1210:
1206:
1202:
1198:
1191:
1189:
1187:
1183:
1178:
1174:
1170:
1166:
1162:
1158:
1151:
1148:
1144:
1138:
1135:
1130:
1126:
1121:
1116:
1112:
1108:
1104:
1100:
1096:
1092:
1088:
1081:
1078:
1073:
1069:
1064:
1059:
1055:
1051:
1047:
1043:
1039:
1035:
1031:
1024:
1021:
1016:
1012:
1008:
1004:
1000:
996:
992:
988:
981:
978:
973:
969:
965:
961:
957:
953:
949:
945:
941:
937:
930:
927:
922:
918:
914:
910:
906:
902:
898:
894:
890:
886:
879:
876:
871:
867:
862:
857:
853:
849:
845:
841:
837:
830:
827:
822:
818:
813:
808:
803:
798:
794:
790:
786:
779:
777:
775:
771:
766:
762:
758:
754:
750:
746:
742:
738:
734:
730:
723:
721:
717:
711:
709:
707:
703:
699:
695:
691:
685:
682:
679:
675:
671:
667:
662:
660:
656:
655:halorhodopsin
652:
647:
645:
641:
637:
633:
632:mental events
629:
625:
621:
620:
615:
611:
607:
603:
599:
595:
591:
587:
583:
579:
575:
567:
565:
562:
558:
554:
548:
546:
545:hyperpolarize
542:
538:
530:
528:
526:
521:
519:
510:
508:
501:
499:
492:
490:
488:
484:
480:
476:
472:
464:
462:
460:
456:
452:
446:
444:
440:
439:
434:
430:
426:
425:
421:
416:
414:
408:
406:
402:
398:
394:
389:
385:
381:
376:
374:
370:
369:
363:
361:
357:
352:
348:
347:
342:
338:
333:
331:
327:
323:
315:
313:
311:
307:
303:
299:
295:
291:
287:
283:
279:
270:
263:
261:
259:
255:
251:
247:
243:
239:
235:
231:
228:
224:
220:
219:
214:
210:
206:
202:
194:
190:
185:
178:
176:
173:
171:
167:
166:
161:
157:
153:
149:
145:
141:
137:
133:
129:
123:
121:
117:
116:
111:
108:
107:Chlorophyceae
103:
101:
96:
89:
87:
85:
81:
77:
73:
72:
67:
63:
59:
56:
55:intracellular
52:
48:
44:
40:
37:
33:
29:
25:
21:
5184:Ion channels
5161:
5070:
5057:Optogenetics
5011:Hegemann lab
4961:(1): 23–33.
4958:
4954:
4914:
4910:
4880:
4876:
4831:
4827:
4817:
4774:
4770:
4760:
4725:
4721:
4711:
4702:
4692:
4665:
4661:
4651:
4618:
4614:
4608:
4576:(1): 23–33.
4573:
4569:
4559:
4527:(1): 23–33.
4524:
4520:
4510:
4467:
4463:
4453:
4408:
4402:
4370:(1): 23–34.
4367:
4363:
4353:
4308:
4304:
4294:
4251:
4247:
4237:
4202:
4198:
4188:
4145:
4141:
4131:
4090:
4086:
4080:
4035:
4031:
4021:
3978:
3974:
3964:
3921:
3917:
3907:
3856:
3852:
3842:
3797:
3793:
3737:
3733:
3677:
3673:
3617:
3613:
3603:
3578:
3572:
3529:
3525:
3515:
3482:
3478:
3468:
3431:
3427:
3417:
3380:
3376:
3366:
3331:
3327:
3317:
3285:(1): 23–34.
3282:
3278:
3268:
3233:
3229:
3215:
3180:
3176:
3166:
3131:
3127:
3117:
3072:
3068:
3058:
3013:
3009:
2999:
2954:
2950:
2940:
2907:
2903:
2897:
2852:
2848:
2838:
2793:
2789:
2779:
2736:
2732:
2722:
2677:
2673:
2663:
2628:
2624:
2578:
2574:
2532:
2528:
2474:
2470:
2460:
2417:
2413:
2403:
2350:
2346:
2336:
2293:
2289:
2279:
2236:
2232:
2222:
2179:
2175:
2125:
2121:
2111:
2078:
2074:
2032:
2028:
1976:
1972:
1962:
1930:(1): 19–25.
1927:
1923:
1875:
1871:
1865:
1838:
1834:
1824:
1789:
1785:
1727:
1723:
1713:
1680:
1676:
1616:
1612:
1602:
1577:
1564:
1521:
1517:
1507:
1464:
1460:
1450:
1407:
1403:
1365:
1361:
1355:
1312:
1308:
1298:
1263:
1259:
1249:
1204:
1200:
1160:
1156:
1150:
1137:
1094:
1090:
1080:
1037:
1033:
1023:
990:
986:
980:
939:
935:
929:
888:
884:
878:
843:
839:
829:
792:
788:
732:
728:
686:
683:
663:
658:
648:
624:Optogenetics
617:
613:
609:
578:transfection
571:
568:Applications
560:
549:
537:depolarizing
534:
524:
522:
514:
505:
496:
471:optogenetics
468:
447:
436:
422:
417:
412:
409:
404:
392:
379:
377:
366:
364:
344:
334:
326:optogenetics
319:
297:
293:
289:
281:
275:
250:metabotropic
216:
198:
174:
168:oocytes and
163:
159:
155:
151:
147:
143:
139:
131:
124:
113:
109:
104:
93:
69:
66:optogenetics
19:
18:
5016:Spudich lab
4883:: 167–189.
4777:(1): 1750.
4254:(1): 4527.
4038:(3): e299.
3859:(1): 7844.
3532:(1): 7844.
651:optogenetic
598:vertebrates
518:motoneurons
397:Roger Tsien
395:. In 2009,
322:genetically
238:Schiff base
213:chromophore
203:. They are
43:green algae
5178:Categories
5006:Boyden lab
712:References
511:Wavelength
475:C-terminal
443:two-photon
356:vertebrate
335:At first,
254:ionotropic
230:derivative
95:Phototaxis
47:phototaxis
28:rhodopsins
5122:Glutamate
4413:CiteSeerX
3881:2041-1723
3740:: 14807.
3712:206554245
3595:237576843
3222:Svoboda K
2932:205404606
2363:CiteSeerX
1641:2041-1723
1619:(1): 65.
1594:253259023
1499:249886382
765:206506942
702:Parkinson
614:correlate
610:recording
602:vitamin A
360:rhodopsin
341:mammalian
234:vitamin A
209:rhodopsin
179:Structure
100:rhodopsin
5021:Kato lab
4985:16600853
4941:17442243
4897:18444900
4858:30396994
4809:29717130
4752:24509078
4703:BBC News
4684:34031601
4635:18432197
4600:16600853
4551:16600853
4502:19151697
4445:13992150
4394:35203089
4345:18697934
4286:34312384
4229:24509633
4180:26113638
4115:17410168
4072:17375185
4032:PLOS ONE
4013:18094685
3956:18682213
3899:36543773
3834:26699459
3772:26443033
3704:24674867
3652:36383037
3564:36543773
3499:21399632
3460:30026684
3409:27011354
3358:24952910
3309:35203089
3260:25762684
3207:24509633
3158:23995068
3109:25201989
3050:21504945
2991:19997631
2951:PLOS ONE
2924:24054067
2889:20543137
2830:19706471
2771:18621808
2714:17483470
2655:21076426
2603:14275254
2595:17435752
2557:17721823
2549:17195846
2509:20473285
2452:20613723
2395:19299587
2328:19389999
2271:17943086
2214:21796121
2152:18432196
2103:15125498
2095:19079251
2049:20081849
2011:19254539
1954:20621963
1900:15096826
1892:16990810
1857:15820685
1816:16360690
1764:16306259
1697:16116447
1659:38167346
1650:10761956
1556:30158696
1491:35726059
1442:26113638
1382:18037436
1347:22266941
1290:18552201
1241:14615590
1177:12565839
1145:AF461397
1129:10049347
821:12060707
757:12089443
706:epilepsy
670:synapses
586:gene gun
541:protists
493:Kinetics
401:kinetics
346:in vitro
343:neurons
264:Function
227:aldehyde
84:protists
5162:Commons
5127:Vesicle
5117:Voltage
5112:Calcium
4976:1459045
4932:3634585
4849:6293277
4800:5931537
4779:Bibcode
4743:3934189
4643:6798764
4591:1459045
4542:1459045
4493:2745650
4472:Bibcode
4385:9758582
4336:2575337
4313:Bibcode
4277:8313717
4256:Bibcode
4220:3943671
4171:4764398
4150:Bibcode
4142:Science
4123:4415339
4095:Bibcode
4063:1808431
4040:Bibcode
4004:3425380
3983:Bibcode
3947:2891506
3926:Bibcode
3890:9772239
3861:Bibcode
3825:4743797
3802:Bibcode
3763:4595828
3742:Bibcode
3682:Bibcode
3674:Science
3643:9733931
3622:Bibcode
3579:bioRxiv
3555:9772239
3534:Bibcode
3507:5907240
3451:6041400
3434:: 451.
3400:4846379
3349:4117813
3300:9758582
3251:4355205
3198:3943671
3149:3793847
3100:4183338
3077:Bibcode
3041:3088623
3018:Bibcode
2982:2780714
2959:Bibcode
2880:2900666
2857:Bibcode
2821:2736443
2798:Bibcode
2762:2553121
2741:Bibcode
2705:1876585
2682:Bibcode
2646:3011044
2500:3177305
2479:Bibcode
2443:3552484
2422:Bibcode
2386:6744370
2355:Bibcode
2347:Science
2319:5262197
2298:Bibcode
2290:Science
2262:6744371
2241:Bibcode
2205:4155501
2184:Bibcode
2143:2692303
2057:7457755
2002:2717302
1981:Bibcode
1945:2995811
1794:Bibcode
1755:1292990
1732:Bibcode
1705:6809511
1621:Bibcode
1578:bioRxiv
1547:6340299
1526:Bibcode
1482:9854242
1433:4764398
1412:Bibcode
1404:Science
1338:4160518
1317:Bibcode
1281:2483371
1209:Bibcode
1120:1300143
1099:Bibcode
1072:8789109
1063:1224992
1042:Bibcode
1015:4309593
995:Bibcode
972:4165365
944:Bibcode
921:4263301
913:6493336
893:Bibcode
870:7010112
737:Bibcode
729:Science
634:, e.g.
594:retinal
574:neurons
438:Cre-lox
424:in vivo
420:rodents
351:retinal
330:neurons
223:retinal
172:(HEK).
165:Xenopus
136:GenBank
90:History
62:calcium
58:acidity
36:sensory
4983:
4973:
4955:Neuron
4939:
4929:
4911:Neuron
4895:
4856:
4846:
4807:
4797:
4750:
4740:
4682:
4641:
4633:
4598:
4588:
4570:Neuron
4549:
4539:
4521:Neuron
4500:
4490:
4464:Nature
4443:
4433:
4415:
4392:
4382:
4343:
4333:
4284:
4274:
4227:
4217:
4178:
4168:
4121:
4113:
4087:Nature
4070:
4060:
4011:
4001:
3975:Nature
3954:
3944:
3897:
3887:
3879:
3832:
3822:
3770:
3760:
3710:
3702:
3650:
3640:
3593:
3562:
3552:
3505:
3497:
3458:
3448:
3407:
3397:
3356:
3346:
3307:
3297:
3258:
3248:
3205:
3195:
3156:
3146:
3107:
3097:
3048:
3038:
2989:
2979:
2930:
2922:
2887:
2877:
2828:
2818:
2769:
2759:
2712:
2702:
2653:
2643:
2601:
2593:
2555:
2547:
2507:
2497:
2471:Nature
2450:
2440:
2414:Nature
2393:
2383:
2365:
2326:
2316:
2269:
2259:
2233:Nature
2212:
2202:
2176:Nature
2150:
2140:
2101:
2093:
2055:
2047:
2009:
1999:
1952:
1942:
1898:
1890:
1855:
1814:
1762:
1752:
1703:
1695:
1657:
1647:
1639:
1592:
1554:
1544:
1518:Nature
1497:
1489:
1479:
1440:
1430:
1380:
1345:
1335:
1309:Nature
1288:
1278:
1239:
1232:283525
1229:
1175:
1127:
1117:
1070:
1060:
1013:
987:Nature
970:
964:628427
962:
936:Nature
919:
911:
885:Nature
868:
861:373196
858:
819:
812:124360
809:
763:
755:
628:causal
619:caused
473:. The
405:
380:
302:cation
298:
292:to 13-
286:photon
160:acop-2
156:acop-1
76:cloned
4639:S2CID
4441:S2CID
4119:S2CID
3708:S2CID
3591:S2CID
3503:S2CID
3377:eLife
2928:S2CID
2599:S2CID
2553:S2CID
2099:S2CID
2053:S2CID
1896:S2CID
1701:S2CID
1590:S2CID
1574:(PDF)
1495:S2CID
1011:S2CID
968:S2CID
917:S2CID
761:S2CID
666:axons
600:) as
588:) or
328:) of
290:trans
282:trans
218:trans
207:like
144:cop-4
140:cop-3
80:algae
4981:PMID
4937:PMID
4893:PMID
4854:PMID
4805:PMID
4748:PMID
4680:PMID
4631:PMID
4596:PMID
4547:PMID
4498:PMID
4431:ISBN
4390:PMID
4341:PMID
4282:PMID
4225:PMID
4176:PMID
4111:PMID
4068:PMID
4009:PMID
3952:PMID
3895:PMID
3877:ISSN
3830:PMID
3768:PMID
3700:PMID
3648:PMID
3560:PMID
3495:PMID
3456:PMID
3405:PMID
3354:PMID
3305:PMID
3256:PMID
3203:PMID
3154:PMID
3105:PMID
3046:PMID
2987:PMID
2920:PMID
2885:PMID
2826:PMID
2767:PMID
2710:PMID
2651:PMID
2591:PMID
2545:PMID
2505:PMID
2448:PMID
2391:PMID
2324:PMID
2267:PMID
2210:PMID
2148:PMID
2091:PMID
2045:PMID
2007:PMID
1950:PMID
1888:PMID
1853:PMID
1835:Cell
1812:PMID
1760:PMID
1693:PMID
1655:PMID
1637:ISSN
1552:PMID
1487:PMID
1438:PMID
1378:PMID
1343:PMID
1286:PMID
1237:PMID
1173:PMID
1125:PMID
1068:PMID
960:PMID
909:PMID
866:PMID
817:PMID
753:PMID
668:and
433:fMRI
386:. A
225:(an
215:all-
193:3ug9
158:and
152:csoB
150:and
148:csoA
142:and
112:and
82:and
4971:PMC
4963:doi
4927:PMC
4919:doi
4885:doi
4844:PMC
4836:doi
4795:PMC
4787:doi
4738:PMC
4730:doi
4726:124
4670:doi
4623:doi
4586:PMC
4578:doi
4537:PMC
4529:doi
4488:PMC
4480:doi
4468:457
4423:doi
4380:PMC
4372:doi
4331:PMC
4321:doi
4309:105
4272:PMC
4264:doi
4215:PMC
4207:doi
4166:PMC
4158:doi
4146:349
4103:doi
4091:446
4058:PMC
4048:doi
3999:PMC
3991:doi
3979:451
3942:PMC
3934:doi
3885:PMC
3869:doi
3820:PMC
3810:doi
3798:113
3758:PMC
3750:doi
3690:doi
3678:344
3638:PMC
3630:doi
3583:doi
3550:PMC
3542:doi
3487:doi
3446:PMC
3436:doi
3395:PMC
3385:doi
3344:PMC
3336:doi
3295:PMC
3287:doi
3246:PMC
3238:doi
3193:PMC
3185:doi
3144:PMC
3136:doi
3095:PMC
3085:doi
3073:111
3036:PMC
3026:doi
3014:108
2977:PMC
2967:doi
2912:doi
2875:PMC
2865:doi
2853:107
2816:PMC
2806:doi
2794:106
2757:PMC
2749:doi
2700:PMC
2690:doi
2678:104
2641:PMC
2633:doi
2583:doi
2537:doi
2495:PMC
2487:doi
2475:465
2438:PMC
2430:doi
2418:466
2381:PMC
2373:doi
2351:324
2314:PMC
2306:doi
2294:324
2257:PMC
2249:doi
2237:450
2200:PMC
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