---
DOAJ_listed: '1'
OA_place: publisher
OA_type: gold
PlanS_conform: '1'
_id: '21473'
abstract:
- lang: eng
text: Physical exercise acutely improves hippocampus-dependent memory. Whereas animal
studies have offered cellular- and synaptic-level accounts of these effects, human
neuroimaging studies show that exercise improves hippocampal-cortical connectivity
at the macroscale level. However, the neurophysiological basis of exercise-induced
effects on hippocampal-cortical circuits remains unknown. Experimental evidence
supports the idea that hippocampal sharp wave-ripples (SWR) play a critical role
in learning and memory. Coupling between SWRs in the hippocampus and neocortex
may reflect modulations in inter-regional connectivity required by mnemonic processes.
Here, we examine the hypothesis that exercise modulates hippocampal-cortical ripple
dynamics in the human brain. We performed intracranial recordings in epilepsy
patients undergoing pre-surgical evaluation, during awake resting state, before
and after an exercise session. Exercise increased ripple rate in the hippocampus.
Exercise also enhanced the coupling and phase-synchrony between cortical ripples
in the limbic and the default mode (DM) cortical networks and hippocampal SWRs.
Further, a higher heart rate during exercise, reflecting exercise intensity, was
related to a subsequent increase in resting state ripples across specific cortical
networks, including the DM network. These results offer the first direct evidence
that a single exercise session elicits changes in ripple events, a well-established
neurophysiological marker of mnemonic processing. The characterisation and anatomical
distribution of the described modulation points to hippocampal ripples as a potential
mechanism by which exercise elicits its reported short-term effects in cognition.
acknowledgement: We acknowledge the generosity of the patients, who contributed time
and effort to take part in this study.
article_number: fcag041
article_processing_charge: Yes
article_type: original
author:
- first_name: Araceli R.
full_name: Cardenas, Araceli R.
last_name: Cardenas
- first_name: Juan F
full_name: Ramirez Villegas, Juan F
id: 44B06F76-F248-11E8-B48F-1D18A9856A87
last_name: Ramirez Villegas
- first_name: Christopher K.
full_name: Kovach, Christopher K.
last_name: Kovach
- first_name: Phillip E.
full_name: Gander, Phillip E.
last_name: Gander
- first_name: Rachel C.
full_name: Cole, Rachel C.
last_name: Cole
- first_name: Andrew J.
full_name: Grossbach, Andrew J.
last_name: Grossbach
- first_name: Hiroto
full_name: Kawasaki, Hiroto
last_name: Kawasaki
- first_name: Jeremy D.W.
full_name: Greenlee, Jeremy D.W.
last_name: Greenlee
- first_name: Matthew A.
full_name: Howard, Matthew A.
last_name: Howard
- first_name: Kirill V.
full_name: Nourski, Kirill V.
last_name: Nourski
- first_name: Matthew I.
full_name: Banks, Matthew I.
last_name: Banks
- first_name: Michelle W.
full_name: Voss, Michelle W.
last_name: Voss
citation:
ama: Cardenas AR, Ramirez Villegas JF, Kovach CK, et al. Exercise enhances hippocampal-cortical
ripple interactions in the human brain. Brain Communications. 2026;8(2).
doi:10.1093/braincomms/fcag041
apa: Cardenas, A. R., Ramirez Villegas, J. F., Kovach, C. K., Gander, P. E., Cole,
R. C., Grossbach, A. J., … Voss, M. W. (2026). Exercise enhances hippocampal-cortical
ripple interactions in the human brain. Brain Communications. Oxford University
Press. https://doi.org/10.1093/braincomms/fcag041
chicago: Cardenas, Araceli R., Juan F Ramirez Villegas, Christopher K. Kovach, Phillip
E. Gander, Rachel C. Cole, Andrew J. Grossbach, Hiroto Kawasaki, et al. “Exercise
Enhances Hippocampal-Cortical Ripple Interactions in the Human Brain.” Brain
Communications. Oxford University Press, 2026. https://doi.org/10.1093/braincomms/fcag041.
ieee: A. R. Cardenas et al., “Exercise enhances hippocampal-cortical ripple
interactions in the human brain,” Brain Communications, vol. 8, no. 2.
Oxford University Press, 2026.
ista: Cardenas AR, Ramirez Villegas JF, Kovach CK, Gander PE, Cole RC, Grossbach
AJ, Kawasaki H, Greenlee JDW, Howard MA, Nourski KV, Banks MI, Voss MW. 2026.
Exercise enhances hippocampal-cortical ripple interactions in the human brain.
Brain Communications. 8(2), fcag041.
mla: Cardenas, Araceli R., et al. “Exercise Enhances Hippocampal-Cortical Ripple
Interactions in the Human Brain.” Brain Communications, vol. 8, no. 2,
fcag041, Oxford University Press, 2026, doi:10.1093/braincomms/fcag041.
short: A.R. Cardenas, J.F. Ramirez Villegas, C.K. Kovach, P.E. Gander, R.C. Cole,
A.J. Grossbach, H. Kawasaki, J.D.W. Greenlee, M.A. Howard, K.V. Nourski, M.I.
Banks, M.W. Voss, Brain Communications 8 (2026).
corr_author: '1'
date_created: 2026-03-22T23:04:34Z
date_published: 2026-03-09T00:00:00Z
date_updated: 2026-03-23T14:30:47Z
day: '09'
ddc:
- '570'
department:
- _id: JoCs
doi: 10.1093/braincomms/fcag041
file:
- access_level: open_access
checksum: b5b45c16defeaf88056fc3b939bd0350
content_type: application/pdf
creator: dernst
date_created: 2026-03-23T14:27:39Z
date_updated: 2026-03-23T14:27:39Z
file_id: '21478'
file_name: 2026_BrainCommunications_Cardenas.pdf
file_size: 33974419
relation: main_file
success: 1
file_date_updated: 2026-03-23T14:27:39Z
has_accepted_license: '1'
intvolume: ' 8'
issue: '2'
language:
- iso: eng
month: '03'
oa: 1
oa_version: Published Version
publication: Brain Communications
publication_identifier:
eissn:
- 2632-1297
publication_status: published
publisher: Oxford University Press
quality_controlled: '1'
scopus_import: '1'
status: public
title: Exercise enhances hippocampal-cortical ripple interactions in the human brain
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 8
year: '2026'
...
---
_id: '12862'
abstract:
- lang: eng
text: Despite the considerable progress of in vivo neural recording techniques,
inferring the biophysical mechanisms underlying large scale coordination of brain
activity from neural data remains challenging. One obstacle is the difficulty
to link high dimensional functional connectivity measures to mechanistic models
of network activity. We address this issue by investigating spike-field coupling
(SFC) measurements, which quantify the synchronization between, on the one hand,
the action potentials produced by neurons, and on the other hand mesoscopic “field”
signals, reflecting subthreshold activities at possibly multiple recording sites.
As the number of recording sites gets large, the amount of pairwise SFC measurements
becomes overwhelmingly challenging to interpret. We develop Generalized Phase
Locking Analysis (GPLA) as an interpretable dimensionality reduction of this multivariate
SFC. GPLA describes the dominant coupling between field activity and neural ensembles
across space and frequencies. We show that GPLA features are biophysically interpretable
when used in conjunction with appropriate network models, such that we can identify
the influence of underlying circuit properties on these features. We demonstrate
the statistical benefits and interpretability of this approach in various computational
models and Utah array recordings. The results suggest that GPLA, used jointly
with biophysical modeling, can help uncover the contribution of recurrent microcircuits
to the spatio-temporal dynamics observed in multi-channel experimental recordings.
acknowledgement: "We thank Britni Crocker for help with preprocessing of the data
and spike sorting; Joachim Werner and Michael Schnabel for their excellent IT support;
Andreas Tolias for help with the initial implantation’s of the Utah arrays.\r\nAll
authors were supported by the Max Planck Society. M.B. was supported by the German\r\nFederal
Ministry of Education and Research (BMBF) through the funding scheme received by\r\nthe
Tübingen AI Center, FKZ: 01IS18039B. N.K.L. and V.K. acknowledge the support from
the\r\nShanghai Municipal Science and Technology Major Project (Grant No. 2019SHZDZX02).
The funders had no role in study design, data collection and analysis, decision
to publish, or preparation of the manuscript. "
article_number: e1010983
article_processing_charge: No
article_type: original
author:
- first_name: Shervin
full_name: Safavi, Shervin
last_name: Safavi
- first_name: Theofanis I.
full_name: Panagiotaropoulos, Theofanis I.
last_name: Panagiotaropoulos
- first_name: Vishal
full_name: Kapoor, Vishal
last_name: Kapoor
- first_name: Juan F
full_name: Ramirez Villegas, Juan F
id: 44B06F76-F248-11E8-B48F-1D18A9856A87
last_name: Ramirez Villegas
- first_name: Nikos K.
full_name: Logothetis, Nikos K.
last_name: Logothetis
- first_name: Michel
full_name: Besserve, Michel
last_name: Besserve
citation:
ama: Safavi S, Panagiotaropoulos TI, Kapoor V, Ramirez Villegas JF, Logothetis NK,
Besserve M. Uncovering the organization of neural circuits with Generalized Phase
Locking Analysis. PLoS Computational Biology. 2023;19(4). doi:10.1371/journal.pcbi.1010983
apa: Safavi, S., Panagiotaropoulos, T. I., Kapoor, V., Ramirez Villegas, J. F.,
Logothetis, N. K., & Besserve, M. (2023). Uncovering the organization of neural
circuits with Generalized Phase Locking Analysis. PLoS Computational Biology.
Public Library of Science. https://doi.org/10.1371/journal.pcbi.1010983
chicago: Safavi, Shervin, Theofanis I. Panagiotaropoulos, Vishal Kapoor, Juan F
Ramirez Villegas, Nikos K. Logothetis, and Michel Besserve. “Uncovering the Organization
of Neural Circuits with Generalized Phase Locking Analysis.” PLoS Computational
Biology. Public Library of Science, 2023. https://doi.org/10.1371/journal.pcbi.1010983.
ieee: S. Safavi, T. I. Panagiotaropoulos, V. Kapoor, J. F. Ramirez Villegas, N.
K. Logothetis, and M. Besserve, “Uncovering the organization of neural circuits
with Generalized Phase Locking Analysis,” PLoS Computational Biology, vol.
19, no. 4. Public Library of Science, 2023.
ista: Safavi S, Panagiotaropoulos TI, Kapoor V, Ramirez Villegas JF, Logothetis
NK, Besserve M. 2023. Uncovering the organization of neural circuits with Generalized
Phase Locking Analysis. PLoS Computational Biology. 19(4), e1010983.
mla: Safavi, Shervin, et al. “Uncovering the Organization of Neural Circuits with
Generalized Phase Locking Analysis.” PLoS Computational Biology, vol. 19,
no. 4, e1010983, Public Library of Science, 2023, doi:10.1371/journal.pcbi.1010983.
short: S. Safavi, T.I. Panagiotaropoulos, V. Kapoor, J.F. Ramirez Villegas, N.K.
Logothetis, M. Besserve, PLoS Computational Biology 19 (2023).
date_created: 2023-04-23T22:01:03Z
date_published: 2023-04-01T00:00:00Z
date_updated: 2025-04-23T08:54:49Z
day: '01'
ddc:
- '570'
department:
- _id: JoCs
doi: 10.1371/journal.pcbi.1010983
external_id:
isi:
- '000962668700002'
pmid:
- '37011110'
file:
- access_level: open_access
checksum: edeb9d09f3e41ba7c0251308b9e372e7
content_type: application/pdf
creator: dernst
date_created: 2023-04-25T08:59:18Z
date_updated: 2023-04-25T08:59:18Z
file_id: '12867'
file_name: 2023_PLoSCompBio_Safavi.pdf
file_size: 4737671
relation: main_file
success: 1
file_date_updated: 2023-04-25T08:59:18Z
has_accepted_license: '1'
intvolume: ' 19'
isi: 1
issue: '4'
language:
- iso: eng
month: '04'
oa: 1
oa_version: Published Version
pmid: 1
publication: PLoS Computational Biology
publication_identifier:
eissn:
- 1553-7358
publication_status: published
publisher: Public Library of Science
quality_controlled: '1'
related_material:
link:
- relation: software
url: https://github.com/shervinsafavi/gpla.git
scopus_import: '1'
status: public
title: Uncovering the organization of neural circuits with Generalized Phase Locking
Analysis
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 19
year: '2023'
...
---
_id: '12149'
abstract:
- lang: eng
text: Editorial on the Research Topic
acknowledgement: This work was supported by a DFG grant ZA990/1 to DZ. This work was
supported by the MSCA EU proposal 841301 - DREAM, European Commission; Horizon 2020
- Research and Innovation Framework Programme to JFRV.
article_number: '1028154'
article_processing_charge: No
article_type: letter_note
author:
- first_name: Giuditta
full_name: Gambino, Giuditta
last_name: Gambino
- first_name: Rebecca
full_name: Bhik-Ghanie, Rebecca
last_name: Bhik-Ghanie
- first_name: Giuseppe
full_name: Giglia, Giuseppe
last_name: Giglia
- first_name: M. Victoria
full_name: Puig, M. Victoria
last_name: Puig
- first_name: Juan F
full_name: Ramirez Villegas, Juan F
id: 44B06F76-F248-11E8-B48F-1D18A9856A87
last_name: Ramirez Villegas
- first_name: Daniel
full_name: Zaldivar, Daniel
last_name: Zaldivar
citation:
ama: 'Gambino G, Bhik-Ghanie R, Giglia G, Puig MV, Ramirez Villegas JF, Zaldivar
D. Editorial: Neuromodulatory ascending systems: Their influence at the microscopic
and macroscopic levels. Frontiers in Neural Circuits. 2022;16. doi:10.3389/fncir.2022.1028154'
apa: 'Gambino, G., Bhik-Ghanie, R., Giglia, G., Puig, M. V., Ramirez Villegas, J.
F., & Zaldivar, D. (2022). Editorial: Neuromodulatory ascending systems: Their
influence at the microscopic and macroscopic levels. Frontiers in Neural Circuits.
Frontiers Media. https://doi.org/10.3389/fncir.2022.1028154'
chicago: 'Gambino, Giuditta, Rebecca Bhik-Ghanie, Giuseppe Giglia, M. Victoria Puig,
Juan F Ramirez Villegas, and Daniel Zaldivar. “Editorial: Neuromodulatory Ascending
Systems: Their Influence at the Microscopic and Macroscopic Levels.” Frontiers
in Neural Circuits. Frontiers Media, 2022. https://doi.org/10.3389/fncir.2022.1028154.'
ieee: 'G. Gambino, R. Bhik-Ghanie, G. Giglia, M. V. Puig, J. F. Ramirez Villegas,
and D. Zaldivar, “Editorial: Neuromodulatory ascending systems: Their influence
at the microscopic and macroscopic levels,” Frontiers in Neural Circuits,
vol. 16. Frontiers Media, 2022.'
ista: 'Gambino G, Bhik-Ghanie R, Giglia G, Puig MV, Ramirez Villegas JF, Zaldivar
D. 2022. Editorial: Neuromodulatory ascending systems: Their influence at the
microscopic and macroscopic levels. Frontiers in Neural Circuits. 16, 1028154.'
mla: 'Gambino, Giuditta, et al. “Editorial: Neuromodulatory Ascending Systems: Their
Influence at the Microscopic and Macroscopic Levels.” Frontiers in Neural Circuits,
vol. 16, 1028154, Frontiers Media, 2022, doi:10.3389/fncir.2022.1028154.'
short: G. Gambino, R. Bhik-Ghanie, G. Giglia, M.V. Puig, J.F. Ramirez Villegas,
D. Zaldivar, Frontiers in Neural Circuits 16 (2022).
date_created: 2023-01-12T12:07:39Z
date_published: 2022-10-26T00:00:00Z
date_updated: 2025-06-12T06:19:09Z
day: '26'
ddc:
- '570'
department:
- _id: JoCs
doi: 10.3389/fncir.2022.1028154
ec_funded: 1
external_id:
isi:
- '000886671400001'
pmid:
- '36405671'
file:
- access_level: open_access
checksum: 457aa00e1800847abb340853058531de
content_type: application/pdf
creator: dernst
date_created: 2023-01-24T10:10:43Z
date_updated: 2023-01-24T10:10:43Z
file_id: '12357'
file_name: 2022_FrontiersNeuralCircuits_Gambino.pdf
file_size: 110031
relation: main_file
success: 1
file_date_updated: 2023-01-24T10:10:43Z
has_accepted_license: '1'
intvolume: ' 16'
isi: 1
keyword:
- Cellular and Molecular Neuroscience
- Cognitive Neuroscience
- Sensory Systems
- Neuroscience (miscellaneous)
language:
- iso: eng
month: '10'
oa: 1
oa_version: Published Version
pmid: 1
project:
- _id: 26BAE2E4-B435-11E9-9278-68D0E5697425
call_identifier: H2020
grant_number: '841301'
name: 'The Brainstem-Hippocampus Network Uncovered: Dynamics, Reactivation and Memory
Consolidation'
publication: Frontiers in Neural Circuits
publication_identifier:
issn:
- 1662-5110
publication_status: published
publisher: Frontiers Media
quality_controlled: '1'
scopus_import: '1'
status: public
title: 'Editorial: Neuromodulatory ascending systems: Their influence at the microscopic
and macroscopic levels'
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 16
year: '2022'
...
---
_id: '8818'
abstract:
- lang: eng
text: The hippocampus has a major role in encoding and consolidating long-term memories,
and undergoes plastic changes during sleep1. These changes require precise homeostatic
control by subcortical neuromodulatory structures2. The underlying mechanisms
of this phenomenon, however, remain unknown. Here, using multi-structure recordings
in macaque monkeys, we show that the brainstem transiently modulates hippocampal
network events through phasic pontine waves known as pontogeniculooccipital waves
(PGO waves). Two physiologically distinct types of PGO wave appear to occur sequentially,
selectively influencing high-frequency ripples and low-frequency theta events,
respectively. The two types of PGO wave are associated with opposite hippocampal
spike-field coupling, prompting periods of high neural synchrony of neural populations
during periods of ripple and theta instances. The coupling between PGO waves and
ripples, classically associated with distinct sleep stages, supports the notion
that a global coordination mechanism of hippocampal sleep dynamics by cholinergic
pontine transients may promote systems and synaptic memory consolidation as well
as synaptic homeostasis.
acknowledgement: We thank O. Eschenko and M. Constantinou for providing feedback on
earlier versions of this work, and J. Werner and M. Schnabel for technical support
during the development of this study. This research was supported by the Max Planck
Society.
article_processing_charge: No
article_type: original
author:
- first_name: Juan F
full_name: Ramirez Villegas, Juan F
id: 44B06F76-F248-11E8-B48F-1D18A9856A87
last_name: Ramirez Villegas
- first_name: Michel
full_name: Besserve, Michel
last_name: Besserve
- first_name: Yusuke
full_name: Murayama, Yusuke
last_name: Murayama
- first_name: Henry C.
full_name: Evrard, Henry C.
last_name: Evrard
- first_name: Axel
full_name: Oeltermann, Axel
last_name: Oeltermann
- first_name: Nikos K.
full_name: Logothetis, Nikos K.
last_name: Logothetis
citation:
ama: Ramirez Villegas JF, Besserve M, Murayama Y, Evrard HC, Oeltermann A, Logothetis
NK. Coupling of hippocampal theta and ripples with pontogeniculooccipital waves.
Nature. 2021;589(7840):96-102. doi:10.1038/s41586-020-2914-4
apa: Ramirez Villegas, J. F., Besserve, M., Murayama, Y., Evrard, H. C., Oeltermann,
A., & Logothetis, N. K. (2021). Coupling of hippocampal theta and ripples
with pontogeniculooccipital waves. Nature. Springer Nature. https://doi.org/10.1038/s41586-020-2914-4
chicago: Ramirez Villegas, Juan F, Michel Besserve, Yusuke Murayama, Henry C. Evrard,
Axel Oeltermann, and Nikos K. Logothetis. “Coupling of Hippocampal Theta and Ripples
with Pontogeniculooccipital Waves.” Nature. Springer Nature, 2021. https://doi.org/10.1038/s41586-020-2914-4.
ieee: J. F. Ramirez Villegas, M. Besserve, Y. Murayama, H. C. Evrard, A. Oeltermann,
and N. K. Logothetis, “Coupling of hippocampal theta and ripples with pontogeniculooccipital
waves,” Nature, vol. 589, no. 7840. Springer Nature, pp. 96–102, 2021.
ista: Ramirez Villegas JF, Besserve M, Murayama Y, Evrard HC, Oeltermann A, Logothetis
NK. 2021. Coupling of hippocampal theta and ripples with pontogeniculooccipital
waves. Nature. 589(7840), 96–102.
mla: Ramirez Villegas, Juan F., et al. “Coupling of Hippocampal Theta and Ripples
with Pontogeniculooccipital Waves.” Nature, vol. 589, no. 7840, Springer
Nature, 2021, pp. 96–102, doi:10.1038/s41586-020-2914-4.
short: J.F. Ramirez Villegas, M. Besserve, Y. Murayama, H.C. Evrard, A. Oeltermann,
N.K. Logothetis, Nature 589 (2021) 96–102.
date_created: 2020-11-29T23:01:19Z
date_published: 2021-01-07T00:00:00Z
date_updated: 2025-07-10T12:01:26Z
day: '07'
department:
- _id: JoCs
doi: 10.1038/s41586-020-2914-4
external_id:
isi:
- '000591047800005'
pmid:
- '33208951'
intvolume: ' 589'
isi: 1
issue: '7840'
language:
- iso: eng
month: '01'
oa_version: None
page: 96-102
pmid: 1
publication: Nature
publication_identifier:
eissn:
- 1476-4687
issn:
- 0028-0836
publication_status: published
publisher: Springer Nature
quality_controlled: '1'
related_material:
link:
- relation: erratum
url: https://doi.org/10.1038/s41586-020-03068-9
scopus_import: '1'
status: public
title: Coupling of hippocampal theta and ripples with pontogeniculooccipital waves
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 589
year: '2021'
...