Abstract
The present work aims to review the structural organization of the mammalian superior colliculus (SC), the putative pathways connecting the SC and the basal ganglia, and their role in organizing complex behavioral output. First, we review how the complex intrinsic connections between the SC’s laminae projections allow for the construction of spatially aligned, visual-multisensory maps of the surrounding environment. Moreover, we present a summary of the sensory-motor inputs of the SC, including a description of the integration of multi-sensory inputs relevant to behavioral control. We further examine the major descending outputs toward the brainstem and spinal cord. As the central piece of this review, we provide a thorough analysis covering the putative interactions between the SC and the basal ganglia. To this end, we explore the diverse thalamic routes by which information from the SC may reach the striatum, including the pathways through the lateral posterior, parafascicular, and rostral intralaminar thalamic nuclei. We also examine the interactions between the SC and subthalamic nucleus, representing an additional pathway for the tectal modulation of the basal ganglia. Moreover, we discuss how information from the SC might also be relayed to the basal ganglia through midbrain tectonigral and tectotegmental projections directed at the substantia nigra compacta and ventrotegmental area, respectively, influencing the dopaminergic outflow to the dorsal and ventral striatum. We highlight the vast interplay between the SC and the basal ganglia and raise several missing points that warrant being addressed in future studies.
Graphical Abstract
[http://dx.doi.org/10.1002/0471733849]
[http://dx.doi.org/10.1016/j.neuron.2021.01.013] [PMID: 33548173]
[http://dx.doi.org/10.1002/cne.901160206] [PMID: 13682733]
[http://dx.doi.org/10.1007/PL00005690] [PMID: 9224849]
[http://dx.doi.org/10.1007/BF00231145] [PMID: 6519226]
[http://dx.doi.org/10.1016/j.neuroimage.2017.12.080] [PMID: 29292132]
[http://dx.doi.org/10.1017/S0952523800000304] [PMID: 2134846]
[http://dx.doi.org/10.1002/cne.901700203] [PMID: 62764]
[http://dx.doi.org/10.1016/j.exer.2005.07.002] [PMID: 16125175]
[http://dx.doi.org/10.1159/000125468] [PMID: 5522341]
[http://dx.doi.org/10.1152/jn.1972.35.4.560] [PMID: 4624740]
[http://dx.doi.org/10.1152/jn.1976.39.4.745] [PMID: 823303]
[http://dx.doi.org/10.1097/00001756-199705260-00001] [PMID: 9223054]
[http://dx.doi.org/10.1152/jn.1945.8.2.123]
[http://dx.doi.org/10.1016/S0079-6123(08)60462-8]
[http://dx.doi.org/10.1016/0165-3806(82)90139-0] [PMID: 6179578]
[http://dx.doi.org/10.1016/0306-4522(84)90007-1] [PMID: 6483194]
[http://dx.doi.org/10.1523/JNEUROSCI.3375-14.2015] [PMID: 26203147]
[http://dx.doi.org/10.1146/annurev-neuro-061010-113728] [PMID: 21456962]
[http://dx.doi.org/10.1523/JNEUROSCI.3137-13.2014] [PMID: 24828636]
[http://dx.doi.org/10.1016/j.neuron.2018.10.021] [PMID: 30392799]
[http://dx.doi.org/10.1523/JNEUROSCI.18-20-08496.1998] [PMID: 9763492]
[http://dx.doi.org/10.1146/annurev.ne.03.030180.001201] [PMID: 6774653]
[http://dx.doi.org/10.1523/JNEUROSCI.3004-13.2014] [PMID: 24647954]
[http://dx.doi.org/10.1113/JP276370] [PMID: 30206945]
[http://dx.doi.org/10.3389/fnana.2017.00036] [PMID: 28487639]
[http://dx.doi.org/10.1016/j.expneurol.2005.08.022] [PMID: 16303128]
[http://dx.doi.org/10.1007/s002219900280] [PMID: 10759167]
[http://dx.doi.org/10.1016/0306-4522(84)90063-0] [PMID: 6462457]
[http://dx.doi.org/10.1002/cne.903190406] [PMID: 1619044]
[http://dx.doi.org/10.1016/0306-4522(86)90267-8] [PMID: 3774146]
[http://dx.doi.org/10.1016/0006-8993(87)90206-X] [PMID: 3607496]
[http://dx.doi.org/10.1007/BF02259139] [PMID: 1541354]
[http://dx.doi.org/10.1007/BF00250606] [PMID: 2466683]
[http://dx.doi.org/10.1038/s41467-021-24241-2] [PMID: 34183678]
[http://dx.doi.org/10.1007/BF00255241] [PMID: 2826204]
[http://dx.doi.org/10.1016/j.cub.2021.04.001] [PMID: 34102128]
[http://dx.doi.org/10.1152/jn.1987.57.1.22] [PMID: 3559673]
[http://dx.doi.org/10.1002/cne.23980] [PMID: 26850989]
[http://dx.doi.org/10.1113/jphysiol.1977.sp011971] [PMID: 903907]
[http://dx.doi.org/10.1152/jn.1975.38.3.690] [PMID: 1127462]
[http://dx.doi.org/10.1523/JNEUROSCI.02-09-01177.1982] [PMID: 7119872]
[http://dx.doi.org/10.1038/299248a0] [PMID: 7110344]
[http://dx.doi.org/10.1152/jn.1983.49.3.674] [PMID: 6834093]
[http://dx.doi.org/10.1016/S0306-4522(01)00600-5] [PMID: 11983321]
[http://dx.doi.org/10.1162/089892902760807230] [PMID: 12495529]
[http://dx.doi.org/10.1152/jn.00369.2003] [PMID: 14534263]
[http://dx.doi.org/10.1152/jn.2001.85.2.506] [PMID: 11160489]
[http://dx.doi.org/10.1152/jn.00639.2003] [PMID: 14973313]
[http://dx.doi.org/10.1159/000121641] [PMID: 7326575]
[http://dx.doi.org/10.1152/jn.00340.2010] [PMID: 20610783]
[http://dx.doi.org/10.1152/jn.90478.2008] [PMID: 18562549]
[http://dx.doi.org/10.1152/jn.1994.72.6.2648] [PMID: 7897481]
[http://dx.doi.org/10.1007/BF00340479] [PMID: 3803481]
[http://dx.doi.org/10.1016/0306-4522(88)90341-7] [PMID: 3362348]
[http://dx.doi.org/10.1152/jn.2002.88.4.2000] [PMID: 12364524]
[http://dx.doi.org/10.1523/JNEUROSCI.06-03-00723.1986] [PMID: 3958791]
[http://dx.doi.org/10.3389/fnsys.2015.00109] [PMID: 26283933]
[http://dx.doi.org/10.1016/0006-8993(86)91080-2] [PMID: 3779395]
[http://dx.doi.org/10.1016/j.neuron.2021.10.004]
[http://dx.doi.org/10.1016/j.conb.2016.06.017] [PMID: 27420378]
[http://dx.doi.org/10.1016/j.neuroscience.2010.12.009] [PMID: 21163336]
[http://dx.doi.org/10.1016/j.neuroscience.2009.10.004] [PMID: 19825395]
[http://dx.doi.org/10.1016/S0166-4328(02)00197-3] [PMID: 12493627]
[http://dx.doi.org/10.1016/j.neuroscience.2004.10.020] [PMID: 15653000]
[http://dx.doi.org/10.1038/nn.4285] [PMID: 27043290]
[http://dx.doi.org/10.1007/BF00464579] [PMID: 6812150]
[http://dx.doi.org/10.1007/BF00250607] [PMID: 3234506]
[http://dx.doi.org/10.1016/j.cub.2018.02.005] [PMID: 29502952]
[http://dx.doi.org/10.1016/0306-4522(88)90345-4] [PMID: 2896313]
[http://dx.doi.org/10.1016/S0304-3940(99)00985-4] [PMID: 10696808]
[http://dx.doi.org/10.1523/ENEURO.0271-20.2020] [PMID: 32928881]
[http://dx.doi.org/10.1016/j.neuroscience.2005.11.015] [PMID: 16361067]
[http://dx.doi.org/10.1016/j.tins.2005.06.006] [PMID: 15982753]
[http://dx.doi.org/10.1016/0306-4522(92)90448-B] [PMID: 1279464]
[http://dx.doi.org/10.3389/fnana.2010.00132] [PMID: 20941324]
[http://dx.doi.org/10.1002/cne.902260306] [PMID: 6747028]
[http://dx.doi.org/10.1007/s12264-022-00858-1] [PMID: 35484472]
[http://dx.doi.org/10.1016/S0079-6123(05)51011-2] [PMID: 16221594]
[http://dx.doi.org/10.3389/fnana.2012.00009] [PMID: 22514521]
[http://dx.doi.org/10.1002/ar.a.20359] [PMID: 16850432]
[http://dx.doi.org/10.1152/jn.00227.2016] [PMID: 27169509]
[http://dx.doi.org/10.1002/cne.903040210] [PMID: 1707899]
[http://dx.doi.org/10.1002/cne.903240308] [PMID: 1401268]
[http://dx.doi.org/10.1002/cne.901870307] [PMID: 114555]
[http://dx.doi.org/10.1016/0006-8993(79)91090-4] [PMID: 90538]
[http://dx.doi.org/10.1126/science.aaa8694] [PMID: 26113723]
[http://dx.doi.org/10.1523/JNEUROSCI.0645-16.2016] [PMID: 27581453]
[http://dx.doi.org/10.1523/JNEUROSCI.2768-14.2014] [PMID: 25274823]
[http://dx.doi.org/10.1523/JNEUROSCI.1493-13.2013] [PMID: 23986231]
[http://dx.doi.org/10.1152/jn.1990.64.2.509] [PMID: 2213129]
[http://dx.doi.org/10.1016/0166-2236(92)90169-9] [PMID: 1377424]
[http://dx.doi.org/10.1016/0006-8993(75)90388-1] [PMID: 1148825]
[http://dx.doi.org/10.1002/cne.903320206] [PMID: 8331213]
[http://dx.doi.org/10.1017/S0952523800008464] [PMID: 7544610]
[http://dx.doi.org/10.1152/jn.01051.2004] [PMID: 15601732]
[http://dx.doi.org/10.1146/annurev-vision-102016-061234] [PMID: 28617660]
[http://dx.doi.org/10.1002/cne.903150209] [PMID: 1372013]
[http://dx.doi.org/10.1152/jn.00705.2003] [PMID: 15010497]
[http://dx.doi.org/10.1002/cne.902830106] [PMID: 2732361]
[http://dx.doi.org/10.1016/0006-8993(84)90050-7] [PMID: 6529626]
[http://dx.doi.org/10.1152/jn.1988.60.1.232] [PMID: 3404219]
[http://dx.doi.org/10.1017/S095252380001261X] [PMID: 9278994]
[http://dx.doi.org/10.3389/fncir.2018.00035]
[http://dx.doi.org/10.1073/pnas.0701934104] [PMID: 17420449]
[http://dx.doi.org/10.1073/pnas.2012937118] [PMID: 33526665]
[http://dx.doi.org/10.1038/s42003-021-02248-1] [PMID: 34117346]
[http://dx.doi.org/10.1523/JNEUROSCI.2305-10.2011] [PMID: 21307233]
[http://dx.doi.org/10.1073/pnas.1504866112] [PMID: 25825743]
[http://dx.doi.org/10.1002/cne.903020111] [PMID: 2086611]
[http://dx.doi.org/10.1046/j.1460-9568.2000.00132.x] [PMID: 10947814]
[http://dx.doi.org/10.1016/j.cell.2014.02.023] [PMID: 24581503]
[http://dx.doi.org/10.1002/syn.10279] [PMID: 14579424]
[http://dx.doi.org/10.1093/cercor/bhy173] [PMID: 30085040]
[http://dx.doi.org/10.1016/0306-4522(85)90199-X] [PMID: 2869444]
[http://dx.doi.org/10.1016/0361-9230(91)90184-L] [PMID: 2015507]
[http://dx.doi.org/10.1126/science.6867718] [PMID: 6867718]
[http://dx.doi.org/10.1152/jn.1996.76.2.1246] [PMID: 8871234]
[http://dx.doi.org/10.1002/cne.901870103] [PMID: 226566]
[http://dx.doi.org/10.1038/nature05860] [PMID: 17522629]
[http://dx.doi.org/10.1016/j.bbr.2007.08.021] [PMID: 17889943]
[http://dx.doi.org/10.1016/j.neuron.2012.02.037] [PMID: 22578499]
[http://dx.doi.org/10.1523/JNEUROSCI.1202-15.2016] [PMID: 26758824]
[http://dx.doi.org/10.1038/nature18601] [PMID: 27357796]
[http://dx.doi.org/10.1038/s41583-020-0292-4] [PMID: 32269316]
[http://dx.doi.org/10.1002/cne.903480103] [PMID: 7814684]
[http://dx.doi.org/10.1111/ejn.15794] [PMID: 35971965]
[http://dx.doi.org/10.1038/nrn3301] [PMID: 22850830]
[http://dx.doi.org/10.1016/j.cobeha.2018.01.017]
[http://dx.doi.org/10.1073/pnas.0900939106] [PMID: 19273843]
[http://dx.doi.org/10.3389/fnsys.2014.00168] [PMID: 25309351]
[http://dx.doi.org/10.1016/j.celrep.2019.01.078]
[http://dx.doi.org/10.1016/j.neuron.2022.06.026] [PMID: 35896109]
[http://dx.doi.org/10.1038/nn.4644] [PMID: 28920934]
[http://dx.doi.org/10.1098/rspb.2022.0799] [PMID: 35703050]
[http://dx.doi.org/10.1073/pnas.1305581110] [PMID: 23918394]
[http://dx.doi.org/10.1073/pnas.89.21.10089] [PMID: 1279669]
[http://dx.doi.org/10.1159/000116574] [PMID: 3334904]
[http://dx.doi.org/10.1016/0166-2236(89)90052-0] [PMID: 2470171]
[http://dx.doi.org/10.1016/j.conb.2008.11.004] [PMID: 19059772]
[http://dx.doi.org/10.1016/j.neuron.2008.09.019] [PMID: 18940594]
[http://dx.doi.org/10.1016/j.bbr.2013.04.040] [PMID: 23643689]
[http://dx.doi.org/10.1126/science.171.3966.82] [PMID: 4992313]
[http://dx.doi.org/10.1113/jphysiol.1980.sp013167] [PMID: 6770082]
[http://dx.doi.org/10.1016/j.cub.2019.07.083] [PMID: 31474533]
[http://dx.doi.org/10.1016/S0306-4522(97)00191-7] [PMID: 9330360]
[http://dx.doi.org/10.1016/0306-4522(88)90344-2] [PMID: 2896312]
[http://dx.doi.org/10.1016/0306-4522(82)90161-0] [PMID: 7078726]
[http://dx.doi.org/10.1016/S0166-2236(84)80197-6]
[http://dx.doi.org/10.1007/BF00236845] [PMID: 3093259]
[http://dx.doi.org/10.1523/JNEUROSCI.0247-09.2009] [PMID: 19403836]
[http://dx.doi.org/10.1038/nn1113] [PMID: 12925855]
[http://dx.doi.org/10.1111/j.1460-9568.2008.06596.x] [PMID: 19175405]
[http://dx.doi.org/10.1038/s41586-018-0078-2] [PMID: 29720647]
[http://dx.doi.org/10.1152/jn.00740.2003] [PMID: 14573557]
[http://dx.doi.org/10.1016/j.neuroscience.2017.07.036] [PMID: 28743454]
[http://dx.doi.org/10.1016/0006-8993(94)91776-0] [PMID: 8180831]
[http://dx.doi.org/10.1016/0006-8993(79)91001-1] [PMID: 314832]
[http://dx.doi.org/10.1016/0361-9230(82)90145-9] [PMID: 6816390]
[http://dx.doi.org/10.1017/S0952523817000050] [PMID: 28965504]
[http://dx.doi.org/10.1002/1096-9861(20000724)423:2<243:AID-CNE5>3.0.CO;2-5] [PMID: 10867657]
[http://dx.doi.org/10.1016/j.cub.2019.10.017]
[http://dx.doi.org/10.1046/j.0953-816x.2001.01712.x] [PMID: 11576195]
[http://dx.doi.org/10.1007/BF00235934] [PMID: 4054280]
[http://dx.doi.org/10.1152/jn.1983.49.5.1230] [PMID: 6864248]
[http://dx.doi.org/10.1152/jn.1989.61.4.780] [PMID: 2723720]
[http://dx.doi.org/10.1007/BF00227139] [PMID: 8224072]
[http://dx.doi.org/10.1016/0006-8993(85)90213-6] [PMID: 2859912]
[http://dx.doi.org/10.1196/annals.1390.012] [PMID: 17360800]
[http://dx.doi.org/10.1038/ncomms7756]
[http://dx.doi.org/10.3389/fnana.2010.00143] [PMID: 21120139]
[http://dx.doi.org/10.1523/JNEUROSCI.1279-17.2017] [PMID: 29175956]
[http://dx.doi.org/10.1016/j.neuron.2016.11.045] [PMID: 27989459]
[http://dx.doi.org/10.1002/(SICI)1096-9861(19990927)412:3<383:AID-CNE2>3.0.CO;2-5] [PMID: 10441229]
[http://dx.doi.org/10.1038/s41467-018-03580-7] [PMID: 29581428]
[http://dx.doi.org/10.1038/s41593-020-00712-5] [PMID: 32989293]
[http://dx.doi.org/10.1016/j.neuron.2019.02.035]
[http://dx.doi.org/10.1523/JNEUROSCI.1606-15.2015] [PMID: 26109669]
[http://dx.doi.org/10.1007/s00429-017-1534-8] [PMID: 28988338]
[http://dx.doi.org/10.1152/jn.00399.2013] [PMID: 24108793]
[http://dx.doi.org/10.1152/jn.00142.2012] [PMID: 22496533]
[http://dx.doi.org/10.1016/j.brainres.2008.01.094] [PMID: 18342841]
[http://dx.doi.org/10.1002/cne.21202] [PMID: 17183537]
[http://dx.doi.org/10.1152/jn.01322.2007] [PMID: 18234976]
[http://dx.doi.org/10.3389/fnsys.2017.00053] [PMID: 28790899]
[http://dx.doi.org/10.1152/jn.01018.2010] [PMID: 21389309]
[http://dx.doi.org/10.1016/j.neuron.2014.07.033] [PMID: 25155959]
[http://dx.doi.org/10.1038/nrn1919] [PMID: 16715055]
[http://dx.doi.org/10.1523/JNEUROSCI.16-10-03444.1996] [PMID: 8627378]
[http://dx.doi.org/10.1007/BF00227239] [PMID: 1397142]
[http://dx.doi.org/10.1152/jn.01009.2002] [PMID: 12611938]
[http://dx.doi.org/10.1038/s41467-020-15443-1] [PMID: 32245963]
[http://dx.doi.org/10.1002/1096-9861(20010305)431:2<198:AID-CNE1065>3.0.CO;2-8] [PMID: 11170000]
[http://dx.doi.org/10.1016/0006-8993(86)90925-X] [PMID: 3730874]
[http://dx.doi.org/10.3389/fnbeh.2014.00115] [PMID: 24765070]
[http://dx.doi.org/10.1016/S0165-0173(02)00181-9] [PMID: 12423763]
[http://dx.doi.org/10.3389/fnbeh.2022.964644] [PMID: 36082310]
[http://dx.doi.org/10.1002/cne.902990206] [PMID: 2172326]
[http://dx.doi.org/10.1002/cne.20342] [PMID: 15514932]
[http://dx.doi.org/10.1002/(SICI)1096-9861(19960122)364:4<637:AID-CNE3>3.0.CO;2-4] [PMID: 8821451]
[http://dx.doi.org/10.1002/cne.901770405] [PMID: 624792]
[http://dx.doi.org/10.1016/0006-8993(85)90214-8] [PMID: 3995318]
[http://dx.doi.org/10.1111/ejn.13413] [PMID: 27717088]
[http://dx.doi.org/10.1002/jnr.25046] [PMID: 35355316]
[http://dx.doi.org/10.1126/sciadv.abe9192] [PMID: 33547085]
[http://dx.doi.org/10.1136/jnnp-2011-302158] [PMID: 23100448]
[http://dx.doi.org/10.1523/JNEUROSCI.22-02-00562.2002] [PMID: 11784803]
[http://dx.doi.org/10.1152/jn.00160.2013] [PMID: 23864369]
[http://dx.doi.org/10.1523/JNEUROSCI.0487-08.2008] [PMID: 18614691]
[http://dx.doi.org/10.1016/j.neuron.2020.04.017] [PMID: 32437650]
[http://dx.doi.org/10.1038/nrn4038] [PMID: 26530468]
[http://dx.doi.org/10.1523/JNEUROSCI.1289-17.2017] [PMID: 29038241]
[http://dx.doi.org/10.1016/0006-8993(78)90881-8] [PMID: 209862]
[http://dx.doi.org/10.1016/S0168-0102(02)00027-5] [PMID: 12067746]
[PMID: 35850655]
[http://dx.doi.org/10.1016/j.neuroscience.2013.07.047] [PMID: 23916713]
[http://dx.doi.org/10.1016/j.neuron.2021.03.017]
[http://dx.doi.org/10.1152/jn.1992.67.6.1615] [PMID: 1629767]
[http://dx.doi.org/10.1002/cne.902360103] [PMID: 2414329]
[http://dx.doi.org/10.1016/0006-8993(90)91087-W] [PMID: 2350684]
[http://dx.doi.org/10.1016/j.neulet.2008.05.071] [PMID: 18562098]
[http://dx.doi.org/10.1152/physrev.2000.80.3.953] [PMID: 10893428]
[http://dx.doi.org/10.1016/0306-4522(94)90208-9] [PMID: 8052406]
[http://dx.doi.org/10.1016/j.neuroscience.2016.05.031] [PMID: 27238892]
[http://dx.doi.org/10.1016/j.neuroscience.2009.04.043] [PMID: 19393292]
[http://dx.doi.org/10.1046/j.1460-9568.2003.02415.x] [PMID: 12534966]
[http://dx.doi.org/10.1152/jn.1998.80.1.1] [PMID: 9658025]
[http://dx.doi.org/10.1126/science.275.5306.1593] [PMID: 9054347]
[http://dx.doi.org/10.1523/JNEUROSCI.1278-13.2013] [PMID: 24259575]
[http://dx.doi.org/10.1016/j.cub.2016.06.041] [PMID: 27780050]
[http://dx.doi.org/10.1016/S0960-9822(00)00593-5] [PMID: 10899013]
[http://dx.doi.org/10.1038/nrn2022] [PMID: 17115078]
[http://dx.doi.org/10.1016/0024-3205(85)90448-5] [PMID: 3990520]
[http://dx.doi.org/10.1016/S0166-4328(98)00102-8] [PMID: 10512583]
[http://dx.doi.org/10.1016/S0165-0173(97)00039-8] [PMID: 9495561]
[http://dx.doi.org/10.1016/S0006-8993(97)00265-5] [PMID: 9221945]
[http://dx.doi.org/10.1152/jn.1992.67.1.145] [PMID: 1552316]
[http://dx.doi.org/10.1126/science.1107026] [PMID: 15746431]
[http://dx.doi.org/10.1016/S0306-4522(00)00019-1] [PMID: 10727783]
[http://dx.doi.org/10.1016/S0166-2236(98)01373-3] [PMID: 10203849]
[http://dx.doi.org/10.1016/j.cell.2015.07.015] [PMID: 26232228]
[http://dx.doi.org/10.1152/jn.00329.2013] [PMID: 24225541]
[http://dx.doi.org/10.1038/s41583-019-0189-2] [PMID: 31171839]
[http://dx.doi.org/10.1016/j.brainresrev.2007.10.007] [PMID: 18055018]
[http://dx.doi.org/10.1016/S0140-6736(13)62418-6]
[http://dx.doi.org/10.1002/ana.25696] [PMID: 32030799]
[http://dx.doi.org/10.1016/j.clinph.2013.01.021]
[http://dx.doi.org/10.1016/bs.pbr.2020.01.001]
[http://dx.doi.org/10.1016/S1474-4422(21)00377-X]
[http://dx.doi.org/10.1016/j.conb.2010.08.008]
[http://dx.doi.org/10.1038/nrneurol.2012.273]
[http://dx.doi.org/10.1093/brain/117.6.1423]
[http://dx.doi.org/10.1016/j.clinph.2022.07.505]
[http://dx.doi.org/10.1152/jn.00094.2007]
[http://dx.doi.org/10.1016/j.neuropsychologia.2009.11.006]
[http://dx.doi.org/10.1038/s41598-020-61572-4]
[http://dx.doi.org/10.1002/acn3.51027]
[http://dx.doi.org/10.1007/s00415-019-09486-8]
[http://dx.doi.org/10.1177/10738584211011979]
[http://dx.doi.org/10.1002/mds.27861]
[http://dx.doi.org/10.1016/j.mehy.2020.110048]