Anatomically segregated basal ganglia pathways allow parallel behavioral modulation


In the basal ganglia (BG), anatomically segregated and topographically organized feedforward circuits are thought to modulate multiple behaviors in parallel. Although topographically arranged BG circuits have been described, the extent to which these relationships are maintained across the BG output nuclei and in downstream targets is unclear. Here, using focal trans-synaptic anterograde tracing, we show that the motor-action-related topographical organization of the striatum is preserved in all BG output nuclei. The topography is also maintained downstream of the BG and in multiple parallel closed loops that provide striatal input. Furthermore, focal activation of two distinct striatal regions induces either licking or turning, consistent with their respective anatomical targets of projection outside of the BG. Our results confirm the parallel model of BG function and suggest that the integration and competition of information relating to different behavior occur largely outside of the BG.

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Fig. 1: Topographically organized projection from BG input to output nuclei.
Fig. 2: Projections of BG output nuclei to downstream targets maintain segregated topography.
Fig. 3: The BG output via Pf forms segregated and closed loops.
Fig. 4: BG output via VM forms segregated closed loops via cortical layer 1.
Fig. 5: Focal optogenetic stimulation of neurons in striatum using TF optics.
Fig. 6: Stimulation of direct pathway projection neurons in VLS induces contralateral licking.
Fig. 7: dSPN stimulation outside of VLS does not interfere with cue-evoked licking.
Fig. 8: DMS and VMS direct pathway stimulation induce contralateral turning.

Data availability

The data that support the findings of this study are available from the corresponding author upon reasonable request.

Code availability

The code used for analysis (MATLAB) is also available from the corresponding author upon reasonable request. Detailed information about software and methods used is available online in the Nature Research Reporting Summary.


  1. 1.

    Roseberry, T. K. et al. Cell-type-specific control of brainstem locomotor circuits by basal ganglia. Cell 164, 526–537 (2016).

    CAS  PubMed  PubMed Central  Google Scholar 

  2. 2.

    Kravitz, A. V. et al. Regulation of parkinsonian motor behaviours by optogenetic control of basal ganglia circuitry. Nature 466, 622–626 (2010).

    CAS  PubMed  PubMed Central  Google Scholar 

  3. 3.

    Tai, L.-H., Lee, A. M., Benavidez, N., Bonci, A. & Wilbrecht, L. Transient stimulation of distinct subpopulations of striatal neurons mimics changes in action value. Nat. Neurosci. 15, 1281–1289 (2012).

    CAS  PubMed  PubMed Central  Google Scholar 

  4. 4.

    Xiong, Q., Znamenskiy, P. & Zador, A. M. Selective corticostriatal plasticity during acquisition of an auditory discrimination task. Nature 521, 348–351 (2015).

    CAS  PubMed  PubMed Central  Google Scholar 

  5. 5.

    Znamenskiy, P. & Zador, A. M. Corticostriatal neurons in auditory cortex drive decisions during auditory discrimination. Nature 497, 482–485 (2013).

    CAS  PubMed  PubMed Central  Google Scholar 

  6. 6.

    Reynolds, J. N. J., Hyland, B. I. & Wickens, J. R. A cellular mechanism of reward-related learning. Nature 413, 67–70 (2001).

    CAS  PubMed  Google Scholar 

  7. 7.

    Shen, W., Flajolet, M., Greengard, P. & Surmeier, D. J. Dichotomous dopaminergic control of striatal synaptic plasticity. Science 321, 848–851 (2008).

    CAS  PubMed  PubMed Central  Google Scholar 

  8. 8.

    Alexander, G. E., DeLong, M. R. & Strick, P. L. Parallel organization of functionally segregated circuits linking basal ganglia and cortex. Annu. Rev. Neurosci. 9, 357–381 (1986).

    CAS  PubMed  Google Scholar 

  9. 9.

    Deniau, J. M. & Chevalier, G. The lamellar organization of the rat substantia nigra pars reticulata: distribution of projection neurons. Neuroscience 46, 361–377 (1992).

    CAS  PubMed  Google Scholar 

  10. 10.

    Deniau, J. M., Menetrey, A. & Charpier, S. The lamellar organization of the rat substantia nigra pars reticulata: segregated patterns of striatal afferents and relationship to the topography of corticostriatal projections. Neuroscience 73, 761–781 (1996).

    CAS  PubMed  Google Scholar 

  11. 11.

    Kolomiets, B. P. et al. Segregation and convergence of information flow through the cortico-subthalamic pathways. J. Neurosci. 21, 5764–5772 (2001).

    CAS  PubMed  PubMed Central  Google Scholar 

  12. 12.

    Kolomiets, B. P., Deniau, J. M., Glowinski, J. & Thierry, A. M. Basal ganglia and processing of cortical information: functional interactions between trans-striatal and trans-subthalamic circuits in the substantia nigra pars reticulata. Neuroscience 117, 931–938 (2003).

    CAS  PubMed  Google Scholar 

  13. 13.

    Kravitz, A. V., Tye, L. D. & Kreitzer, A. C. Distinct roles for direct and indirect pathway striatal neurons in reinforcement. Nat. Neurosci. 15, 816–818 (2012).

    CAS  PubMed  PubMed Central  Google Scholar 

  14. 14.

    Yttri, E. A. & Dudman, J. T. Opponent and bidirectional control of movement velocity in the basal ganglia. Nature 533, 402–406 (2016).

    CAS  PubMed  PubMed Central  Google Scholar 

  15. 15.

    Wang, L., Rangarajan, K. V., Gerfen, C. R. & Krauzlis, R. J. Activation of striatal neurons causes a perceptual decision bias during visual change detection in mice. Neuron 97, 1369–1381.e5 (2018).

    CAS  PubMed  PubMed Central  Google Scholar 

  16. 16.

    Kelly, R. M. & Strick, P. L. Macro-architecture of basal ganglia loops with the cerebral cortex: use of rabies virus to reveal multisynaptic circuits. Prog. Brain Res. 143, 447–459 (2004).

  17. 17.

    Saunders, A. et al. A direct GABAergic output from the basal ganglia to frontal cortex. Nature 521, 85–89 (2015).

    CAS  PubMed  PubMed Central  Google Scholar 

  18. 18.

    Jackson, A. & Crossman, A. R. Subthalamic nucleus efferent projection to the cerebral cortex. Neuroscience 6, 2367–2377 (1981).

    CAS  PubMed  Google Scholar 

  19. 19.

    Aoki, S. et al. An open cortico-basal ganglia loop allows limbic control over motor output via the nigrothalamic pathway. eLife 8, e49995 (2019).

    PubMed  PubMed Central  Google Scholar 

  20. 20.

    Sippy, T., Lapray, D., Crochet, S. & Petersen, C. C. H. Cell-type-specific sensorimotor processing in striatal projection neurons during goal-directed behavior. Neuron 88, 298–305 (2015).

    CAS  PubMed  PubMed Central  Google Scholar 

  21. 21.

    Babl, S. S., Rummell, B. P. & Sigurdsson, T. The spatial extent of optogenetic silencing in transgenic mice expressing channelrhodopsin in inhibitory interneurons. Cell Rep. 29, 1381–1395.e4 (2019).

    CAS  PubMed  Google Scholar 

  22. 22.

    Li, N. et al. Spatiotemporal constraints on optogenetic inactivation in cortical circuits. eLife 8, e48622 (2019).

    CAS  PubMed  PubMed Central  Google Scholar 

  23. 23.

    Travers, J. B., Dinardo, L. A. & Karimnamazi, H. Motor and premotor mechanisms of licking. Neurosci. Biobehav. Rev. 21, 631–647 (1997).

    CAS  PubMed  Google Scholar 

  24. 24.

    Wang, S. & Redgrave, P. Microinjections of muscimol into lateral superior colliculus disrupt orienting and oral movements in the formalin model of pain. Neuroscience 81, 967–988 (1997).

    CAS  PubMed  Google Scholar 

  25. 25.

    Sahibzada, N., Dean, P. & Redgrave, P. Movements resembling orientation or avoidance elicited by electrical stimulation of the superior colliculus in rats. J. Neurosci. 6, 723–733 (1986).

    CAS  PubMed  PubMed Central  Google Scholar 

  26. 26.

    Capelli, P., Pivetta, C., Esposito, M. S. & Arber, S. Locomotor speed control circuits in the caudal brainstem. Nature 551, 373–377 (2017).

    CAS  PubMed  Google Scholar 

  27. 27.

    Zingg, B. et al. AAV-mediated anterograde transsynaptic tagging: mapping corticocollicular input-defined neural pathways for defense behaviors. Neuron 93, 33–47 (2017).

    CAS  PubMed  Google Scholar 

  28. 28.

    Zingg, B., Peng, B., Huang, J., Tao, H. W. & Zhang, L. I. Synaptic specificity and application of anterograde transsynaptic AAV for probing neural circuitry. J. Neurosci. 40, 3250–3267 (2020).

    CAS  PubMed  PubMed Central  Google Scholar 

  29. 29.

    Hunnicutt, B. J. et al. A comprehensive excitatory input map of the striatum reveals novel functional organization. eLife 5, e19103 (2016).

    PubMed  PubMed Central  Google Scholar 

  30. 30.

    Hintiryan, H. et al. The mouse cortico-striatal projectome. Nat. Neurosci. 19, 1100–1114 (2016).

    CAS  PubMed  PubMed Central  Google Scholar 

  31. 31.

    Hooks, B. M. et al. Topographic precision in sensory and motor corticostriatal projections varies across cell type and cortical area. Nat. Commun. 9, 1–16 (2018).

    Google Scholar 

  32. 32.

    Mastro, K. J., Bouchard, R. S., Holt, H. A. K. & Gittis, A. H. Transgenic mouse lines subdivide external segment of the globus pallidus (GPe) neurons and reveal distinct GPe output pathways. J. Neurosci. 34, 2087–2099 (2014).

    CAS  PubMed  PubMed Central  Google Scholar 

  33. 33.

    Mana, S. & Chevalier, G. The fine organization of nigro-collicular channels with additional observations of their relationships with acetylcholinesterase in the rat. Neuroscience 106, 357–374 (2001).

    CAS  PubMed  Google Scholar 

  34. 34.

    Mandelbaum, G. et al. Distinct cortical-thalamic-striatal circuits through the parafascicular nucleus. Neuron 102, 636–652.e7 (2019).

    CAS  PubMed  PubMed Central  Google Scholar 

  35. 35.

    Klapoetke, N. C. et al. Independent optical excitation of distinct neural populations. Nat. Methods 11, 338–346 (2014).

    CAS  PubMed  PubMed Central  Google Scholar 

  36. 36.

    Kuramoto, E. et al. Ventral medial nucleus neurons send thalamocortical afferents more widely and more preferentially to layer 1 than neurons of the ventral anterior–ventral lateral nuclear complex in the rat. Cereb. Cortex 25, 221–235 (2015).

    PubMed  Google Scholar 

  37. 37.

    Rubio-Garrido, P., Pérez-de-Manzo, F., Porrero, C., Galazo, M. J. & Clascá, F. Thalamic input to distal apical dendrites in neocortical layer 1 is massive and highly convergent. Cereb. Cortex 19, 2380–2395 (2009).

    PubMed  Google Scholar 

  38. 38.

    Krout, K. E., Loewy, A. D., Westby, G. W. M. & Redgrave, P. Superior colliculus projections to midline and intralaminar thalamic nuclei of the rat. J. Comp. Neurol. 431, 198–216 (2001).

    CAS  PubMed  Google Scholar 

  39. 39.

    McHaffie, J. G., Stanford, T. R., Stein, B. E., Coizet, V. & Redgrave, P. Subcortical loops through the basal ganglia. Trends Neurosci. 28, 401–407 (2005).

    CAS  PubMed  Google Scholar 

  40. 40.

    Chevalier, G. & Deniau, J. M. Spatio-temporal organization of a branched tecto-spinal/ tecto-diencephalic neuronal system. Neuroscience 12, 427–439 (1984).

    CAS  PubMed  Google Scholar 

  41. 41.

    Oh, S. W. et al. A mesoscale connectome of the mouse brain. Nature 508, 207–214 (2014).

    CAS  PubMed  PubMed Central  Google Scholar 

  42. 42.

    Rossi, M. A. et al. A GABAergic nigrotectal pathway for coordination of drinking behavior. Nat. Neurosci. 19, 742–748 (2016).

    CAS  PubMed  PubMed Central  Google Scholar 

  43. 43.

    Evans, D. A. et al. A synaptic threshold mechanism for computing escape decisions. Nature 558, 590–594 (2018).

    CAS  PubMed  PubMed Central  Google Scholar 

  44. 44.

    Shang, C. et al. A parvalbumin-positive excitatory visual pathway to trigger fear responses in mice. Science 348, 1472–1477 (2015).

    CAS  PubMed  Google Scholar 

  45. 45.

    Pisano, F. et al. Depth-resolved fiber photometry with a single tapered optical fiber implant. Nat. Methods 16, 1185–1192 (2019).

    CAS  PubMed  Google Scholar 

  46. 46.

    Pisanello, M. et al. Tailoring light delivery for optogenetics by modal demultiplexing in tapered optical fibers. Sci. Rep. 8, 4467 (2018).

  47. 47.

    Pisanello, F. et al. Dynamic illumination of spatially restricted or large brain volumes via a single tapered optical fiber. Nat. Neurosci. 20, 1180–1188 (2017).

    CAS  PubMed  PubMed Central  Google Scholar 

  48. 48.

    Gong, S. et al. Targeting Cre recombinase to specific neuron populations with bacterial artificial chromosome constructs. J. Neurosci. 27, 9817–9823 (2007).

    CAS  PubMed  PubMed Central  Google Scholar 

  49. 49.

    Tecuapetla, F., Matias, S., Dugue, G. P., Mainen, Z. F. & Costa, R. M. Balanced activity in basal ganglia projection pathways is critical for contraversive movements. Nat. Commun. 5, 4315 (2014).

  50. 50.

    Yin, H. H. & Knowlton, B. J. The role of the basal ganglia in habit formation. Nat. Rev. Neurosci. 7, 464–476 (2006).

    CAS  PubMed  Google Scholar 

  51. 51.

    Guo, Z. V. et al. Flow of cortical activity underlying a tactile decision in mice. Neuron 81, 179–194 (2014).

    CAS  PubMed  Google Scholar 

  52. 52.

    Cebrián, C., Parent, A. & Prensa, L. Patterns of axonal branching of neurons of the substantia nigra pars reticulata and pars lateralis in the rat. J. Comp. Neurol. 492, 349–369 (2005).

    PubMed  Google Scholar 

  53. 53.

    Oldenburg, I. A. & Sabatini, B. L. Antagonistic but not symmetric regulation of primary motor cortex by basal ganglia direct and indirect pathways. Neuron 86, 1174–1181 (2015).

    CAS  PubMed  PubMed Central  Google Scholar 

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We thank members of the Sabatini laboratory, W. Regehr, M. Andermann and N. Uchida for discussions. We thank J. Levasseur for mouse husbandry and genotyping, and J. Saulnier and L. Worth for laboratory administration. We thank W. Kuwamoto, J. Grande, M. Ambrosino, B. Pryor and E. Lubbers for assistance with behavioral experiments and histology. This work was supported by the NIH (grant no. NINDS R01NS103226), a P30 Core Center Grant (grant no. NINDS NS072030), an Iljou Foundation scholarship and a grant from the Simons Collaborative on the Global Brain.

Author information




J.L. and B.L.S. conceptualized the study, wrote the original draft, and reviewed and edited the manuscript. J.L. performed and participated in all experiments and W.W. performed the slice electrophysiology experiments. B.L.S. supervised the study and was responsible for acquisition of funding.

Corresponding author

Correspondence to Bernardo L. Sabatini.

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Competing interests

B.L.S. is a founder of and holds private equity in Optogenix. Tapered fibers commercially available from Optogenix were used as tools in the research.

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Peer review information Nature Neuroscience thanks Marc Fuccillo and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

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Extended data

Extended Data Fig. 1 Topography is maintained across the AP axis of SNr.

a, Example histology section in anterior SNr and posterior SNr (see main text and methods for injection protocol). Row represent AP coordinates and each column indicates the location of the AAV1.cre for anterograde tracing (see main text). Scale bar, 500um. Topographical labelling was also observed in anterior part of SNr. Similar results were obtained in n = 9 mice (n = 3 for each site).

Extended Data Fig. 2 EP to LHb projection is topographic.

a, Experiment protocol. AAV1-cre and AAV1-Flpo in medial and lateral striatum respectively, followed by a cocktail of viruses encoding either DIO-Tdtom or fDIO-EYFP in EP. b, left, histological sections showing the anterogradely labelled cells from medial (red) and lateral (green) striatum. right, Axonal fibers innervating LHb in a non-overlapping fashion. Similar results were obtained in n = 2 mice. Scale bar, 1mm, 1mm, 0.1mm from left to right panel.

Extended Data Fig. 3 Topography of SNr output is consistent across mice.

a, Histology sections for all injected mice for experiment in Fig. 2c. First row represents column the injection site, and second to fourth column represent axonal fibers in Pf, VM and SC. Each row represents individual mouse. Scale bar, 250um.

Extended Data Fig. 4 SNr output to SC segregated both across medial-lateral axis and across layers.

a, left, Example histology section with mSNr axons (red) and lSNr axons (green) from experiment described in Fig. 2c. right, Cartoon diagram summarizing the region and layer specificity of lSNr and mSNr observed. lSNr innervated the lateral SC (abbreviations: Zo: Zona layer of the superior colliculus, Op: optic nerve layer of the superior colliculus, InG: Intermediate gray layer of the superior colliculus, InWh: Intermediate white layer of the superior colliculus, DpG: Deep gray layer of the superior colliculus). lSNr innervated the lSC in InWh, and extending to upper layer of InWh in mSC. mSNr innervated the superfifical part of InG and deeper part of InWh in mSC. Similar results were obtained in n = 4 mice. Scale bar, 500um. b, left, Example histological section with VLSSNr (yellow), DLSSNr (cyan), and DMSSNr (magenta) axons in SC (images reproduced from Fig. 2f). right, Carton diagram summarizing the region and layer specificity of VLSSNr, DLSSNr, and DMSSNr observed. VLSSNr targeted the InWh part of lSC, DLSSNr targeted the central part of SC, extending to the upper layer of InWh in mSC, and DMSSNr targeted the upper InG and the lower InWh. Thus, lSNr axons seem to be a combination of VLSSNr and DLSSNr. Scale bar, 500um. Similar results were obtained in n = 9 mice, 3 for each site.

Extended Data Fig. 5 SNr output to Zona Incerta is topographically organized.

a, Example coronal section showing mSNr (red:TdTom)and lSNr (green:EYFP) axon fibers in zona incerta, from experiment described in Fig. 2a. Similar results were obtained in n = 4 mice. b, Example coronal sections showing DMSSNr (magenta:TdTom), DLSSNr (cyan:TdTom) and VLSSNr (yellow:TdTom), axon fibers in zona incerta, from experiment described in Fig. 2e. Similar results were obtained in n = 9 mice, n = 3 for each site.

Extended Data Fig. 6 Brain regions targeted by SNr.

a, Example histology section of brain regions targeted by DMSSNr, DLSSNr, and VLSSNr. Each row represent a brain region, and each column is a different SNr subpopulation labelled from distinct striatal regions (see experiment described in Fig. 2e). b, top left, Quantification of normalized fluorescence intensity across brain regions for DMSSNr, DLSSNr, and VLSSNr (see Methods). bottom right, Similar quantification of brain stem regions, IRt and PCRt, but normalized only to total fluorescence in brain stem. Abbreviations. VM/VA: ventromedial/ventral anterior thalamus, PC/CL/CM: paracentral/centrolateral/central medial thalamus, Pf: parafascicular nucleus, LH: lateral hypothalamus, IC: inferior colliculus, SC: superior colliculus, ZI: zona incerta, PAG: paracqeuductal gray, MA3: medial accessory oculomotor nucleus, MLR: mesencephalic locomotor region, PnO: pontine reticular nucleus, oral part, LDTg: laterodorsal tegmental nucleus, PB: parabrachical nuecleus, IRt: intermediate reticular nucleus, PCRt: parvicellular reticular nucelus, Gi: gigantocellular reticular nucleus (DMSSNr: n=2 mice; DLSSNr: n=3 mice; VLSSNr: n=3 mice).

Extended Data Fig. 7 Anterogradely labelled putative dopamine axons in striatum.

a, Example coronal sections showing putative dopamine axons in striatum (from experiment described in Fig. 2e). Each row represent a distinct striatal injection of AAV1.cre from DMS, DLS and VLS. Injection site is shown in green (H2b-EGFP), dopamine axons in red (TdTom). For all three striatal injections, dopamine axons tend to co-localize with the injection site. Similar results were obtained in n = 9 mice, 3 for each site. b, Schematic showing experiment for validating that the axons observed in striatum are dopamine axons. AAV1-Flpo was injected striatum, followed by a mixture of AAV-DIO-TdTom and AAV-Coff/Fon-EYFP in SNr/SNc, in a Slc6a3-IRES-Cre mouse, where Cre is expressed in dopamine neurons. This allowed us to anterogradely label SNr neurons, similar to experiment in Fig. 2e, but excluding the dopamine neurons (DA-/STR recipient), while also labelling dopamine neurons as a control. c, Example sagittal section showing the injection site. Dopamine neurons are expressing TdTom (red) whereas non-dopaminergic/anterogradely labelled cells are expressing EYFP (green). Similar results were obtained in n = 2 mice. d, Sagittal sections showing axons in striatum (left), thalamus (middle) and SC (right). Dopaminergic axons (red:TdTom) were only seen in striatum, whereas non-dopaminergic/striatal recipient axon (green:EYFP) were only seen in thalamus and SC. Given the lack of EYFP fibers in striatum, axons seen in striatum from anterograde tracing at striatum (experiment in Fig. 2e) likely are dopamine axons. Similar results were obtained in n = 2 mice.

Extended Data Fig. 8 SC projects back to Pf and VM in a topographical fashion.

a, Overlap of SNr axons labelled via anterograde tracing in striatum (AAV1.Cre, see Fig. 2e) and cortical axons. Scale bar, 500um. bottom, from Allen Institute for Brain Science. Allen Mouse Brain Connectivity Atlas (2011). Available from b, Experimental protocol for labelling medial and lateral SC. Two anterograde tracers (AAV1.Cre and AAV1.Flpo) were injected in ACA/tjM1 respectively, followed by AAV.DIO.TdTom or AAV.fDIO.EYFP in medial or lateral SC. This allowed labelling of mSC and lSC without leaking into other regions outside SC. c, Example coronal section showing the injection site in SC (mSC in red, lSC in green). Similar results were obtained in n = 3 mice. Scale bar, 1mm. d, Example coronal sections showing the axonal targets of mSC and lSC. We observed topography in both Pf and VM, similar to SNr topography (see main Text). Similar results were obtained in n = 3 mice. Scale bar, 1mm.

Extended Data Fig. 9 Detailed optical setup for TF stimulation.

The optical setup allows depth dependent optical illumination combined with TF. The main components are: P1: pockel cell (or any power modulator), S1: shutter, M1: piezo mirror, M3: small mirror (0.5” diameter), M2: galvo mirror, L1: Achromatic doublet (AC508-200-A-ML, f=200mm, diameter 2”, Thorlabs), L2: Aspheric condenser (ACL5040-A, f=40mm, diameter 50mm, Thorlabs), X1: XY translation cage mount + Z-axis translation mount (Thorlabs). M2 galvo mirror was used to change the incident angle onto the patchcord attached at the end of X1. M1 was used to correct for any misalignment for each angle.

Extended Data Fig. 10 dSPNs VLS stimulation engage cortical and collicular BG loops.

a, Schematic showing protocol for recording activity in tjM1 and lSC while stimulating VLS dSPNs. Mice were implanted with a single tapered fiber targeting VLS, and injected with AAV-DIO-CoChR in VLS, similar to experiment described in Fig. 6a (see methods). Extracellular recording with a silicon probe was done in tjM1 and lSC, on the same side (right hemisphere) as the stimulation side in striatum (n = 2 mice). b, Mean firing rate during the inter-trial-interval where mice were required to withhold licking, and during which VLS dSPNs stimulation caused contralateral licking (Fig. 6). Mean firing rate of both tjM1 (left) and lSC (right) increased during stimulation (blue, 100 ms stim) relative to no stimulation trial (grey). Shaded light blue represents laser on period (100ms) (n = 102 units in tjM1, n = 65 units in lSC) (mean ± s.e.m across neurons). c, Fraction of cells that were significantly modulated by dSPNs stimulation in VLS in tjM1 (top) and SC (bottom). Cells are categorized into either excited (blue), inhibited (red) or no sigficant change (grey). The majority of cells recorded (53% in tjM1, 68% in lSC) were excited by the stimulation (0-500 ms window relative to laser onset, p<0.05, Mann–Whitney U test, see Methods). d, Mean firing rate, during tone presentation, of cells that were significantly modulated during ITI stimulation. Firing rate for tjM1 (left) and lSC (right) in left trials and right trials. Firing rate during no stim (grey) and during stim trials (blue). Shaded light blue represents laser on period (100ms) (n=102 units in tjM1, n=65 units in lSC) (mean ± s.e.m across neurons).

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Lee, J., Wang, W. & Sabatini, B.L. Anatomically segregated basal ganglia pathways allow parallel behavioral modulation. Nat Neurosci 23, 1388–1398 (2020).

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