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. Author manuscript; available in PMC: 2025 Feb 1.
Published in final edited form as: Neurosci Biobehav Rev. 2023 Dec 22;157:105523. doi: 10.1016/j.neubiorev.2023.105523

Table 1.

The circadian regulation of rodent behaviors.

Task Animal Treatment Experimental Time Intertrial interval Results References
Passive avoidance Swiss Webster mouse ZT time 12:12h LD, ZT9 or 17 24h Light phase groups performed better than dark ones. (Stephens et al., 1967)
Passive avoidance Rat ZT time; melanocyte - stimulating hormone 12:12h LD, ZT6 or 13 48h Light phase groups performed better than dark ones. Melanocytes improved their performance in the dark. (Sandman et al., 1971)
Passive avoidance Sprague-Dawley rat ZT time 12:12h LD, every 4 h 48h Light phase groups performed better than dark ones. (Davies et al., 1973)
Passive avoidance Rat ZT time; ITI 12:12h LD, ZT2, 8, 14, 20 15min, 6h, 12h, 18h, 24h, 30h, 36h, 48h, 72h 15min and all 24h groups performed better than others. (Holloway and Wansley, 1973b)
Passive avoidance Rat ZT time; ITI 12:12h LD, ZT1-4.5 or 5.5-8 30sec-36h 30 sec, 15min, 12h, 24h, and 36h groups performed better than others. (Wansley and Holloway, 1976)
Passive avoidance Sprague-Dawley rat ZT time; ITI 12:12h LD; ZT1-4 or 7-10 15min or every 3-h up to 24h 15min, 9h, 12h, and 24h groups performed better than others. (Hunsicker and Mellgren, 1977)
Passive avoidance Sprague-Dawley rat SCN lesions; ITI 12:12h LD; ZT6-7 18h, 24h, or 30h 24h performance was better than others; SCN lesions did not follow this trend. (Stephan and Kovacevic, 1978)
Passive avoidance Rat +12h, +6h, or −6h shifts 12:12h LD, ZT0.5-2.5 48-h or 7 days Phase-shifted groups had impaired performance. (Tapp and Holloway, 1981)
Passive avoidance Wistar rat +4h or −6h shifts 14:10h LD→DL or DD 24h, 48h, 96h, 144h Phase-shifted groups had impaired performance. (Fekete et al., 1985)
Passive avoidance Wistar rat −6h shifts; ACTH analog ORG-2766 or DGAVP 14:10h LD→DL 24h, 48h,168h Phase-shifted groups had impaired performance; ACTH and DGAVP attenuated the deficits. (Fekete et al., 1986)
Passive avoidance Long-Evans rat ZT time; aging 12:12h DL, ZT13 or 23 24h or 6 weeks In the early dark phase, performance was better than in the late dark phase tested at 6 weeks, but not 24 h, later in the aged, but not young, group (Winocur and Hasher, 1999)
Fear conditioning C57BL/6J mouse ZT time 12:12h LD; ZT2 or 14 Every 24-h up to 5 days Dark phase groups performed better than light ones. (Valentinuzzi et al., 2001)
Fear conditioning (tone-cued) C-57/6J; C-3H; mouse ZT time 12:12h LD or DL or DD, ZT3 or 15 N.A. Light phase groups performed better than dark ones. (Chaudhury and Colwell, 2002)
Fear conditioning C57Bl/6 mouse ZT time and time cues 12:12h LD, ZT4-8 or 16-20 (tested at the same or different times) 24h, 36h, or 48h ZT4 groups had better performances than others. (Eckel-Mahan et al., 2008b)
Fear conditioning (tone-cued) Long-Evans rat Every +3h shift for 6 days(one session) or 4 sessions 12:12h LD, ZT3 24h No effects of phase shifts. (Craig and McDonald, 2008)
Fear conditioning C57Bl/6 mouse Phase shift before or after the LD 12:12h LD, ZT3 or 6 1 to 7days The phase-shifted group had impaired performance. (Loh et al., 2010)
Fear conditioning C57Bl/6 mouse SCN lesions 12:12h LD 2weeks SCN lesions impaired performance. (Phan et al., 2011)
Fear conditioning (extinction) Sprague-Dawley rat ZT time 12:12h LD, ZT4 or 16 24h Facilitated extinction more in the dark phase group than in the light one. (Woodruff et al., 2015b)
Fear conditioning C57BL/6J mouse +12h shift or bifurcated 6:6h LD 12;12h LD, ZT12 or 24 12h or 24h The Phase-shifted group showed impaired retrieval, while the bifurcated group showed impaired learning. (Harrison et al., 2017)
Fear conditioning C57Bl6/J mouse ZT time; ITI; Bmal1 deletion 12:12h LD→DD, CT4 or 16 30min, 42h or 54h The CT4 group had better long-term retrieval than CT16; Bmal1 deletion impaired performance. (Price and Obrietan, 2018)
Fear conditioning (extinction) Sprague-Dawley rat ZT time; Per1/2 knockdown in PFC 12:12h LD, ZT4 or 16 24h Facilitated extinction in the dark phase group; PFC Per1/2 knockdown eliminated this facilitation. (Woodruff et al., 2018)
Fear conditioning (extinction) C57Bl/6J mouse +8h shifted across 1 or 2 days 12:12h LD, ZT13 24h or 48h The phase-shifted group showed impaired fear of extinction. (Clark et al., 2020)
Fear conditioning Wistar rat ZT time; DD 12:12h LD or DD, ZT The DD group showed facilitated fear extinction, specifically during the light phase. (Asadian et al., 2022)
Fear conditioning Siberian hamster −3 h shift 16:8h LD, ZT16-20 48h The phase-shifted group showed impaired fear memory. (Steiger et al., 2022)
MWM Long-Evans rat +3h shift 12:12h LD, ZT11 2 or 10 days (probe) There were no effects on learning; the phase-shifted group showed impaired retention. (Devan et al., 2001)
MWM (delayed non-match-to-sample) Long-Evans rat ZT time and time cues; aging 12:12h DL, ZT13 or 23 0-80sec The early dark phase group performed better than the late dark phase group in the old animals, whereas the reverse was true for the young. (Winocur and Hasher, 2004)
MWM Wistar rat ZT time 12:12h LD, ZT3 or 15 30 min Dark phase groups performed better than light ones. (Valentinuzzi et al., 2004a)
MWM C57BL/6 mouse LL 12:12h LD or LL N.A. (no probe) The LL group showed impaired learning more than the LD one. (Fujioka et al., 2011)
MWM Diurnal grass rat ZT time 12:12h LD, ZT4 or 16 24h There were no effects on learning; the light phase groups performed better than the dark ones. (Martin-Fairey and Nunez, 2014a)
MWM ICR mouse Abnormal LD 12:12h, 3:5h or 5:3h LD N.A. Abnormal LD impaired the performance. (Li et al., 2017)
MWM Wistar Kyoto and SHR rat LD disturbance 12:12h LD, ZT1-2 24h (probe) The LD disturbance impaired the performance. (Wang et al., 2020)
MWM Wistar rat LL or DD; fluoxetine 12:12h LD→ LL or DD, ZT0-3 24h The LL group showed impaired performance, while fluoxetine reversed the deficits. (Sharma et al., 2021)
MWM Long-Evans rat 12:9h LD for 6 days 12:12h LD→12:9h LD, ZT10.5 18 days (probe) The 21-day group showed impaired memory, but not learning. (Deibel et al., 2022)
MWM Wistar rat ZT time; DD 12:12h LD or DD, ZT The DD group showed impaired spatial memory. (Asadian et al., 2022)
MWM Non-Tg and 3xTg-AD mice ZT time; 3xTg-AD 12:12h LD, ZT4 or 16 24h (probe) The light group learned faster than the dark one, while the dark group had better reversal learning than the light one. AD impaired probe and reversal memory. (Carvalho da Silva et al., 2022)
NOR Siberian hamster ZT time; arrhythmic circadian; PTZ (1 mg/kg, i.p. daily) 16:8h LD, ZT3, 7, 11, 15, 19, 23 60min Early light groups had impaired performance in controls; the arrhythmic group showed impaired performance; PTZ facilitated performance (Ruby et al., 2008)
NOR Wistar rat ZT time 12:12h LD, ZT2, 12, 14 or 20 Min or 24h No effects in time-of-day. (Takahashi et al., 2013)
NOR Djungarian hamster ZT time; arrhythmic circadian 14:10h LD, ZT4, 7, 13, 16, 19 1h ZT13, 16, and 19 groups performed better than ZT4 and 7 groups, and arrhythmic circadian impaired performance. (Müller et al., 2015)
NOR Djungarian hamster Arrhythmic circadian 14:10h LD 1h Arrhythmic circadian impaired performance (Weinert et al., 2016)
NOR C57BL/6 mouse ZT time; DD; SON lesions 12:12h LD, ZT0, 4, 8, 12, 16, 20 24h Early light and SCN lesioned groups showed impaired performance. (Shimizu et al., 2016)
NOR & object-displacement recognition C57BL/6J mouse ZT time; LL 12:12h LD, ZT6 or 18; 12:12h LD→LL, CT6 or 18 5min or 24h The light phase group performed better than the dark phase one. (Tam et al., 2017)
NOR B6/C3H mouse Ts65Dn; SCN lesions 12:12h LD, ZT6 24h SCN lesions did not affect memory in controls; Ts65Dn mice had impaired performance, while SCN lesions alleviated it in Ts65Dn mice. (Chuluun et al., 2020)
NOR (one object during the learning and removal in the test) C57BL/6N Crl mouse ZT time 12:12h LD or DD, ZT3.5 or 15.5 N.A. The light phase groups performed better than the dark ones in learning, but not memory. (McCauley et al., 2020)
NOR C57BL/6 mouse ZT time; dim light (ZT12-16) 12:12h LD, ZT2 or 14 12h The light phase group performed better than dark phase one; dim light exposure reversed this pattern. (Tam et al., 2021)
NOR Wistar rat LL or DD; fluoxetine 12:12h LD→ LL or DD, ZT0-3 24h The LL group showed impaired memory, while fluoxetine reversed the deficits. (Sharma et al., 2021)
NOR C57BL/6J mouse Cry1, Cry2 KO, AAV-Cry1 12:12h LD→DD, ZT20-22 24h AAV-Cry1 expression in the SCN attenuated the deficits of Cry1-Cry2 KO. (Maywood et al., 2021)
NOR C57BL/6 mouse ZT time 12:12h LD, ZT0, 6, 12, 18 12h or 24h No effects of time of day (the dark phase groups seemed to perform better in complex object recognition). (Gessner et al., 2022)
NLR C57BL/6J mouse ZT time; sex 12:12h LD→DD, CT4 or 16 30min The dark phase group performed better than the light one in males, while the opposite effect was found in females. (Goode et al., 2022)
Active avoidance Sprague-Dawley rat ZT time; ACTH 12:12h LD, ZT1.5-2.5 or 9-10 Variable ratio: 90 sec The late light phase group performed better than the early light phase one; ACTH facilitated performance (Pagano and Lovely, 1972)
Active avoidance (extinction) Rat ZT time; ITI 12:12h LD, ZT0-4, 7-10, 12-15, 18-21 15min, 6h, 12h, 18h, 24h 15min, 12h, and 24h groups had stronger resistance to extinction than the others. (Holloway and Sturgis, 1976)
Active avoidance (extinction) Rat ZT time; pinealectomy 12;12h LD, ZT3 or 15 60sec The dark phase group learned faster than the light one. (Catalá et al., 1985)
Shock avoidance operant task Sprague-Dawley rat ZT time 12:12h LD or DL, ZT2, 6 or 10, 14, 18, 22 N.A. Dark phase groups performed better in shock avoidance, but not responses, than light ones. (Ghiselli and Patton, 1976)
CPA (foot shock) Syrian hamster ITI 14:10h LD, 24, 48, 32, or 40h 24 and 48h groups were more effective than the 32 and 40h groups. (Cain et al., 2004a)
CPA (foot shock) Syrian hamster ZT time and time cues 14:10h LD, ZT4 or 13 24h Non-matched LD cues impaired performance (Cain et al., 2008)
CPA (foot shock) Syrian hamster ZT time and time cues; SCN lesions 14:10h LD, ZT2 or 11 24h Non-matched LD cues impaired performance: SCN lesions did not affect it. (Cain et al., 2012)
CPP (wheel running) Syrian hamster ZT time 14:10h LD, ZT4 or 13; LL, CT4 or 13 24h The effects were paired with LD cues. (Ralph et al., 2002)
CPP (food) Rat Time cue 12:12h LD, CT2 or 11 (tested at the same or different times) 24h No effects of time cues. (McDonald et al., 2002)
CPP (wheel running) Syrian hamster ZT time; SCN lesions 14:10h LD, ZT3 or 14 24h The effects were paired with LD cues, and SCN was not needed. (Ko et al., 2003)
CPP (food) Wistar and Long-Evans rats ZT time and time cues 12:12h LD, ZT3 or 11 (training at ZT11) 24h No effects were observed in the Long-Evan rats; non-matched LD cues affected the Wistar rats. (Cain et al., 2004b)
CPP Syrian hamster +6h shift every 3 days for 25 days 14:10h LD, ZT18 8 days The phase-shifted group showed impaired performance. (Gibson et al., 2010)
CPP (food) Long-Evans rat Every +3h shift for 6 days→LD 10days, repeated 4 times 12:12h LD 24h The phase-shifted group showed impaired performance. (McDonald et al., 2013)
CPP (morphine) Wistar rat ZT time and time cue 12:12h LD, ZT1-2 or 11-12 24h The late-light phase group showed a stronger preference than the early-light ones. (Khaksari et al., 2022)
Radial-arm maze Sprague-Dawley rat DL; theophylline (15mg/kg, i.p. daily) 12:12h LD or DL 7 or 14 days The dark phase group performed better than the light one; theophylline facilitated performance only in the light phase. (Hauber and Bareiß, 2001)
Radial-arm maze C3H/HeN mouse ZT time; Per1 depletion 12:12h LD, ZT2 or 14 24h The light phase group performed better than the dark ones; Per1 depletion reversed this pattern. (Rawashdeh et al., 2014)
T-maze operant task Wistar rat ZT time; SCN ablation 12:12h LD→DD, ZT3-4, or 10-11 Variable ratio 10 sec Intact learning without LD cues and SCN. (Mistlberger et al., 1996)
Spontaneous T-maze Siberian hamster ZT time; arrhythmic circadian; PTZ (0.3 or 1 mg/kg, i.p. daily) 16:8h LD, ZT3, 7, 11, 15, 19, 23 N.A. Late light and dark phase groups performed better than early light ones; PTZ improved performanc e in the arrhythmic circadian group. (Ruby et al., 2013)
Spontaneous T-maze Siberian hamster Arrhythmic circadian; SCN lesions 16:8h LD, N.A. Arrhythmic circadian lesions, but not SCN lesions, impaired performance. (Fernandez et al., 2014)
Spontaneous T-maze C57BL/6J mouse ZT time; diet 12:12h LD, ZT2 or 14 N.A. The dark phase group performed better than the light one; high fat and sucrose diets impaired it. (Davis et al., 2020)
Spontaneous T-maze C57BL/6J mouse ZT time; diet 12:12h LD, ZT2 or 14 N.A. The dark phase group performed better than the light one; a high-fat diet impaired it. (Davis et al., 2021)
Spontaneous T-maze C57BL/6J mouse ZT time; diet 12:12h LD, ZT2 or 14 N.A. The dark phase group performed better than the light one; a high-fat diet impaired it. (Davis et al., 2021)
Spontaneous Y-maze Wistar rat LL or DD; fluoxetine 12:12h LD→ LL or DD, ZT0-3 N.A. LL group showed impaired memory, while fluoxetine reversed the deficits. (Sharma et al., 2021)
Spontaneous T-maze C57BL/6J mouse ZT time; Tg-SwDI 12:12h LD, ZT2 or 14 N.A. The dark phase group performed better than the light one; the Tg-SwDI mouse had impaired performance. (Fusilier et al., 2021)
6-point alley T-maze C57Bl/6 mouse ZT time 12:12h LD, ZT2, 6, 10, 14, 18, 22 N.A. Dark phase groups performed better than light ones. (Hoffmann and Balschun, 1992)
Spontaneous Y-maze C57BL/6J mouse SCN lesions 12:12h LD N.A. SCN lesions did not affect performance. (Mulder et al., 2014)
Sustained attention operant task Sprague-Dawley rat ZT time 12:12h LD, ZT4 or 16 N.A. The dark phase group performed better than the light one. (Gritton et al., 2012b)
Sustained attention operant task Sprague-Dawley rat ZT time; SCN lesions 12:12h LD, ZT 4 or 16 N.A. The dark phase group performed better than the light one; SCN lesions impaired performance at ZT4. (Gritton et al., 2013)
Barnes maze Mouse ZT time; Bmal1 forebrain deletion 12:12h LD, ZT4 or 16 17days (probe) Bmal1 deletion led to learning and recall deficits at both times. (Price et al., 2016)
Barnes maze C57BL/6J mouse ZT time; elF4E or MNK depletion 12:12h LD, ZT6 or 18 24h (probe) Dark phase groups performed better than light ones; elF4E and MNK depletions affected learning but not memory. (Liu et al., 2022)
Social recognition Djungarian hamster Arrhythmic circadian; social ranking 14:10h LD 2min or 24h The social subordinate, but not dominant, animals had intact 24-h memory; arrhythmic circadian impairment impaired the memory (Weinert et al., 2016)
Social recognition C57BL/6N mouse ZT time; time cues; Bmal1 deletion 12:12h LD, ZT4 or 10 6h or 24h ZT10 showed impaired retrieval of weak social memory in WT; Bmal1 deletion impaired memory at both times. (Hasegawa et al., 2019)
Licking latency task Rat ITI 12:12h LD, ZT1.5-4 or 5.5-8 15min, 1-h or every 6h up to 36h 12h, 24h, and 36h groups performed better than others. (Wansley and Holloway, 1975)
Circadian time-place three-arms task C57BL/6J mouse ZT time; Cry depletion 12:12h LD, ZT2-3.5, 5-6.5 or 8-9.5 3h Animals were able to learn specific time-place rules, but the Cry gene knockout impaired it. (Van der Zee et al., 2008)

Abbreviations: ZT: zeitgeber time; CT: circadian time; LD or DL: light-dark cycle; DD: dark-dark cycle; MWM: Morris water maze; N.A. not applicable or within seconds; CTA: conditioned taste aversion; CPP: conditioned place preference; CPA: conditioned place avoidance; ACTH: adrenocorticotropic hormone; DGAVP: desglycinamide−9-(Arg8)-vasopressin; PTZ: pentylenetetrazol; SHR: spontaneously hypertensive rat; PFC; prefrontal cortex; Tg-SwDI: the human APP gene with K670N/M671L, E693Q and D694N mutations; 3xTg AD: the TauP301L, APPSwe, and γ-secretase (PS1M146V); NOR, novel object recognition with 24 h training-testing interval; TMh, time memory to heat exposure; cTPL, circadian time place memory; CFC, contextual fear conditioning; NLR, novel location recognition; CR, conditioned reflex; Ts65Dn: segmental trisomic 16 on the B6/C3h background