Introduction to our PhD Programme
Understanding how neuronal activity supports brain function and redressing aberrant dynamics when the brain goes awry in diseases requires a trans-disciplinary and cross-species approach, integrating experimental and clinical investigations with engineering and computational frameworks. Our PhD programme at the Medical Research Council Brain Network Dynamics Unit (MRC BNDU) trains a future generation of neuroscientists to combine these diverse techniques in innovative research projects serving both foundational, theoretical and translational neuroscience. Ours is a vibrant community of scientists and students drawn from around the world.
PhD students at the MRC BNDU can choose among diverse yet complementary research projects that leverage from a substantial array of research techniques. They benefit from the guidance of two supervisors and the feedback from the outstanding research community, locally in the Unit and in the wider intellectual environment of the University of Oxford. Students also have opportunities to further their knowledge and skills through seminars, including our Unit “Methods meetings” and practical tutorials.
Techniques
Students apply for one or more of the research projects made available each year. Projects involve the intensive use of a focussed set of trans-disciplinary methods but students also become familiar with the full range of approaches taken within the Unit. These include:
- Behavioural experiment paradigms with rodents and humans
- Advanced signals analysis, including statistical analytical techniques, mathematical modelling and machine learning.
- Cell type-specific monitoring including electron microscopy, optogenetics and fast-scan cyclic voltammetry.
- Computational modelling.
- System and cell type specific brain stimulation using closed-loop approaches and magnetic, electrical, pharmacological and optogenetic stimulation.
In addition, to the above we actively develop novel research tools for neuroscience discovery, and better disease treatments.
ArchT-GFP–expressing neurons in the pyramidal cell layer of mouse hippocampus. From: Recoding a cocaine-place memory engram to a neutral engram in the hippocampus. Trouche S. et al. Nat Neurosci. 2016 Apr;19(4):564-7.
“The Unit has provided three keys ingredients to kickstart my scientific career: a constant source of inspiration, the freedom to explore my ideas, and frequent opportunities for feedback.”
Ben and Flavie demonstrate how electricity in the muscles of the arm can be detected and used to control a robot claw, during a presentation to primary school children.
Career Development
Career Development is central to the Unit’s ethos. Every student at the Unit benefits from all the resources the MRC BNDU offers, as well as those offered by our host department, Nuffield Department of Clinical Neurosciences, the Medical Sciences Division, and the student’s own college. Within the MRC BNDU the Training, Career Development and Capacity Building Strategy Committee meets regularly to oversee training for our students and early career post-docs. Regular events in the Unit include the biannual science day, which offers a chance for the whole unit to come together and share new and exciting results in a collegiate atmosphere. We run an annual training day which is open to all and focusses on different aspects of career development for academic and non-academic careers. Recent sessions have included developing a patient and public involvement strategy, combining a career with caring responsibilities and a workshop on delivering grant pitches. Previous students at the Unit have all gone on to careers in Academia, Medicine, or Industry.
“Previous students who completed their PhD in the Unit have gone on to successful careers in a wide range of job roles. The challenging yet rewarding research in the Unit is a great start to your career and I would definitely recommend working here!”
Projects
Supervisors: Prof Huiling Tan
Applications are invited from both Home students and International students to join a multidisciplinary team of scientists studying neurodynamics and its control for therapy. This studentship is available from the start of academic year 2021/22 for 3 years, and will be primarily supervised by Associate Professor Huiling Tan at the MRC BNDU.
Background: Brain-Computer Interface technology (BCI) aims to decode a user’s mental state or movement intent through brain signals, and use this information to operate a computer interface, wheelchair, robotic arm or other device. BCI is not only used to provide an alternative pathway for the brain to interact with the environment, but also provides a potential tool for neurofeedback training, in which patients may learn to modulate their own brain activities through reinforcement learning. In addition, the BCI framework is also used to deliver stimulation to the brain in a closed-loop fashion based on the detection of specific neural signals or specific mental states in order to improve the treatment of neurological disorders such as Parkinson’s disease (PD).
Project Description: The overall aim of the PhD project is to interface with the brain to modify behaviour and improve therapy. To achieve this goal, we will investigate the function of frequency-specific neural oscillations in decision-making and motor control in both health and disease through behavioural experiments with electrophysiological recordings and controlled neural stimulation in humans. The project will also involve identifying and extracting features in electrophysiological signals in real-time, as well as the design, implementation and test of BCIs for neurofeedback training and/or for closed-loop neuromodulation to interact with brain activities.
Training: The project will take place in the Medical Research Council Brain Network Dynamics Unit at the University of Oxford and students will benefit from the extensive generic and transdisciplinary skills training and career development opportunities available within the Unit. This particular project will also offer the following specific training: 1. Behavioural experiment paradigms with healthy participants and patients with neurological disorders; 2. Advanced signals analysis, including statistical analytical techniques, mathematical modelling and machine learning; 3. Invasive and non-invasive brain stimulation using closed-loop approaches.
Key Publications:
He S, Everest-Phillips C, Clouter A, Brown P and Tan H (2020) Neurofeedback-Linked Suppression of Cortical β Bursts Speeds Up Movement Initiation in Healthy Motor Control: A Double-Blind Sham-Controlled Study. Journal of Neuroscience, 40 (20): 4021-4032
Little S, Brown P (2020) Debugging Adaptive Deep Brain Stimulation for Parkinson's Disease. Movement Disorder, 35(4):555-561.
Tan H, Debarros J, He S, Pogosyan A, Aziz TZ, Huang Y, Wang S, Timmermann L, Visser-Vandewalle V, Pedrosa DJ, Green AL, Brown P (2019) Decoding voluntary movements and postural tremor based on thalamic LFPs as a basis for closed-loop stimulation for essential tremor. Brain Stimulation, 12(4):858-867.
Funding: This Ph.D. (D.Phil.) studentship is funded by the Medical Research Council (MRC), a part of UKRI. The successful applicant is entitled to receive a tax-free stipend and, as a minimum, tuition fees paid at the Home level, regardless of whether they are Home or International students. Please see further details about MRC/UKRI studentships and updated guidance regarding Home and International eligibility. Successful applicants can also be considered for University scholarships.
Qualifications & Experience: Interested candidates should possess, or expect to receive, a 1st class or upper 2nd class degree (or equivalent) in a related scientific discipline, e.g. biological or physical sciences, medicine, computer science, engineering, mathematics. Good analytical and programming skills are essential. Previous experience in brain computer interfacing, electronics or neuroscience research is desired.
Contact: We strongly encourage candidates to contact the project supervisor before submitting an application. To find out more about this MRC-funded studentship, the research project, and the application process, please contact Prof Huiling Tan by email on huiling.tan@ndcn.ox.ac.uk. Prof Tan is also happy to consider supervising PhD students funded by Departmental scholarships and other doctoral training programmes at Oxford, as well as PhD students with their own independent funding.
How to apply: To be considered for this MRC-funded studentship, please submit an application for admission to the D.Phil. in the Nuffield Department of Clinical Neurosciences (course code RD_CU1), following the guidance. On the application form, in the section headed ‘Departmental Studentship Applications’, please indicate that you are applying for a studentship and enter the reference code for a MRC BNDU studentship "21CLINNEURO01WEB".
Supervisor: David Dupret
Applications are invited from both Home students and International students to join a multidisciplinary team of scientists studying neural dynamics of memory circuits in the rodent brain. Up to two MRC-funded studentships are available from the start of academic year 2021/22, for 3 years each, and both will be primarily supervised by Professor David Dupret at the MRC BNDU.
Background: Memory is central to adaptation and survival, enabling individuals to predict from past experience what can happen next in an ever-changing environment. The idea that groups of neurons transiently synchronize their spiking activity to organize information-representing cell assemblies is central to the scientific investigation our laboratory carries at the nexus between brain and behaviour.
Project Description: The overall aim of the PhD project is to investigate how neuronal activities distributed across brain regions allow the division of computational labour involved in complex, multimodal memories that hold information spanning the sensory, motor and cognitive domains. To achieve this goal, we will investigate the concerted action of groups of neurons scattered across brain regions as well as frequency-specific network oscillations during memory tasks. The project will also involve identifying electrophysiological signals in real-time to implement closed-loop neuromodulation of brain activities.
Training: The project will take place in the Medical Research Council Brain Network Dynamics Unit at the University of Oxford and students will benefit from the extensive generic and transdisciplinary skills training and career development opportunities available within the Unit. As a whole, this particular project will expose you to a large array of techniques that are central to the field of Systems Neuroscience. Notably, you will receive advanced training in (1) large-scale electrophysiological recordings of neuronal activity from the rodent brain, (2) optogenetic manipulation of molecularly-defined neuronal populations (with or without closed-loop systems), (3) behavioural assays of learning and memory in rodents, (4) multivariate data analyses (notably using graph-theoretical tools) and programming (Python, R, and Jupyter). We will offer you the possibility to either actively engage with all of these techniques, or concentrate on the analyses of existing electrophysiological datasets that comprise spike trains and local field potentials.
Key Publications:
Barron H.C., Reeve H.M, Koolschijn R.S., Perestenko P.V., Shpektor A., Nili H., R. Rothaermel, Campo-Urriza N., O’Reilly J.X., Bannerman D.M., Behrens T.E.J. and Dupret D. Neuronal computation underlying inferential reasoning in humans and mice. Cell. 2020 Oct 1;183(6):1-16.
Trouche S, Koren V, Doig NM, Ellender TJ, El-Gaby M, Lopes-Dos-Santos V, Reeve HM, Perestenko PV, Garas FN, Magill PJ, Sharott A and Dupret D. A Hippocampus-Accumbens Tripartite Neuronal Motif Guides Appetitive Memory in Space. Cell. 2019 Mar 7;176(6):1393-1406.
Lopes-dos-Santos V, van de Ven GM, Morley A, Trouche S, Campo-Urriza N and Dupret D. Parsing hippocampal theta oscillations by nested spectral components during spatial exploration and memory-guided behavior. Neuron. 2018 Nov 21;100(4):940-952.
van de Ven GM, Trouche S, McNamara CG, Allen K, Dupret D. Hippocampal Offline Reactivation Consolidates Recently Formed Cell Assembly Patterns during Sharp Wave-Ripples. Neuron. 2016 Dec 7;92(5):968-974.
Trouche S, Perestenko PV, van de Ven GM, Bratley CT, McNamara CG, Campo-Urriza N, Black SL, Reijmers LG, Dupret D. Recoding a cocaine-place memory engram to a neutral engram in the hippocampus. Nat Neurosci. 2016 Apr;19(4):564-7.
Funding: These Ph.D. (D.Phil.) studentships are funded by the Medical Research Council (MRC), a part of UKRI. All successful applicants are entitled to receive a tax-free stipend and, as a minimum, tuition fees paid at the Home level, regardless of whether they are Home or International students. Please see further details about MRC/UKRI studentships and updated guidance regarding Home and International eligibility. Successful applicants can also be considered for University scholarships.
Qualifications & Experience: Interested candidates should possess, or expect to receive, a 1st class or upper 2nd class degree (or equivalent) in a related scientific discipline, e.g. biological or physical sciences, medicine, computer science, engineering, mathematics. Previous experience in neuroscience research is essential. This includes clear contribution to either in vivo electrophysiological recordings, behavioural assays of learning and memory in rodents, and/or data analyses and programming (e.g., Python, R, and Jupyter). Note that appointed students will be offered the possibility to either actively engage with all of these techniques, or concentrate on the analyses of existing electrophysiological datasets that comprise spike trains and local field potentials.
Contact: We strongly encourage candidates to contact the project supervisor before submitting an application. To find out more about these MRC-funded studentships, the research project, and the application process, please contact Professor David Dupret by email on david.dupret@bndu.ox.ac.uk. Professor Dupret is also happy to consider supervising PhD students funded by Departmental scholarships and other doctoral training programmes at Oxford, as well as PhD students with their own independent funding.
How to apply: To be considered for one of these MRC-funded studentships, please submit an application for admission to the D.Phil. in the Nuffield Department of Clinical Neurosciences (course code RD_CU1), following the guidance. On the application form, in the section headed ‘Departmental Studentship Applications’, please indicate that you are applying for a studentship and enter the reference code for a MRC BNDU studentship "21CLINNEURO01WEB".
Supervisor: Rafal Bogacz
Applications are invited from both Home students and International students to join a multidisciplinary team of scientists studying computational models of brain decision networks. Up to two MRC-funded studentships are available from the start of academic year 2021/22, for 3 years each, and both will be primarily supervised by Professor Rafal Bogacz at the MRC BNDU.
Background: Neurons releasing dopamine modulate information processing and learning in a brain network playing a key role in selection of movements and more general decision making. Understanding the function of dopamine is important, because the death of dopaminergic neurons in Parkinson’s disease leads to difficulties in movement initiation. A classical reinforcement learning theory (Schultz et al 1997 Science 275, 1593-1599) has provided a beautiful description of dopamine function in learning about rewards, but has not explained why dopamine depletion results in motor impairments. This question was addressed by a recent modelling framework called DopAct (Bogacz 2020 eLife 9, e53262) which has extended the classical theory to also describe dopamine function in planning actions to get the rewards. Although DopAct explains the key data on dopaminergic activity and effects of dopamine depletion, this theory needs to be extended in multiple directions to fully account for experimentally observed patterns of dopamine release and their effects.
Project Description: The goal of the project is to extend the theory of dopamine to describe its effects on the dynamics of various neural populations it modulates, describe the results of dopamine depletion in Parkinson’s disease, and the effects of treatments. The work on the project will involve development and analysis of mathematical models, computer simulations, and comparison with experimental data gathered in MRC BNDU and by other collaborators in Oxford. The applicants need to have strong mathematical skills and background (in calculus, linear algebra and probability theory), and be proficient in computer programming.
Training: The project will take place in the Medical Research Council Brain Network Dynamics Unit at the University of Oxford and students will benefit from the extensive generic and transdisciplinary skills training and career development opportunities available within the Unit. This particular project will also offer specific training in mathematical modelling, computer simulations, data analysis and machine learning.
Funding: These Ph.D. (D.Phil.) studentships are funded by the Medical Research Council (MRC), a part of UKRI. All successful applicants are entitled to receive a tax-free stipend and, as a minimum, tuition fees paid at the Home level, regardless of whether they are Home or International students. Please see further details about MRC/UKRI studentships and updated guidance regarding Home and International eligibility. Successful applicants can also be considered for University scholarships.
Qualifications & Experience: Interested candidates should possess, or expect to receive, a 1st class or upper 2nd class degree (or equivalent) in a related scientific discipline, e.g. computer science, engineering, mathematics, biological or physical sciences. Strong mathematical skills and background (in calculus, linear algebra and probability theory) are essential, as is proficiency in computer programming.
Contact: We strongly encourage candidates to contact the project supervisor before submitting an application. To find out more about these MRC-funded studentships, the research project, and the application process, please contact Professor Rafal Bogacz by email on rafal.bogacz@bndu.ox.ac.uk. Professor Bogacz is also happy to consider supervising PhD students funded by Departmental scholarships and other doctoral training programmes at Oxford, as well as PhD students with their own independent funding.
How to apply: To be considered for one of these MRC-funded studentships, please submit an application for admission to the D.Phil. in the Nuffield Department of Clinical Neurosciences (course code RD_CU1), following the guidance. On the application form, in the section headed ‘Departmental Studentship Applications’, please indicate that you are applying for a studentship and enter the reference code for a MRC BNDU studentship "21CLINNEURO01WEB".
The closing date for applications is 12.00 midday UK time on Friday 8th January 2021.
Interviews for short-listed applicants will be held between 25th and 28th January 2021.
Supervisors: Prof Hayriye Cagnan and Prof Rick Helmich
Background: Tremor is one of the most common symptoms of Parkinson’s disease and could impact approximately 85% of the patient population. However, despite its prevalence our understanding of tremor in Parkinson’s disease is limited, influencing treatment of this common symptom. To date, pharmaceutical therapies available for the treatment of Parkinson’s disease had limited effect on tremor; with some studies reporting approximately 60% of patients’ tremor not responding to dopaminergic medication. These observations have motivated our research into alternative stimulation based treatment strategies for tremor such as peripheral median nerve stimulation, non-invasive transcranial alternating current stimulation and invasive deep brain stimulation. This work has exposed an important consideration to take into account while developing novel treatment strategies: the natural variability of tremor. Natural variability of tremor which has consequences over undulations in tremor severity and how patient’s trembling limb moves in space over seconds, hours and days has a significant impact on how we evaluate efficacy of new treatment strategies. Beyond its impact on treatment development, we also hypothesize that natural variabilities in tremor, as assessed from wearable technologies, provide a window into their origin in the central brain network.
Project Description: We will explore the cerebral mechanisms underlying spontaneous transitions between different tremor modes, by testing for brain activity time-locked to these events using combined tri-axial accelerometery and fMRI based on the Personalised Parkinson’s Project dataset. Finally, by grouping data across patients depending on how often their tremor switches between different modes, we will evaluate potential links between tremor-related brain activity and inter-regional connectivity, using combined accelerometery-fMRI, and measures of their natural tremor variability as evaluated from the wearable sensors.
Training: The project will take place in the Medical Research Council Brain Network Dynamics Unit at the University of Oxford and students will benefit from the extensive generic and transdisciplinary skills training available within the Unit. The student will spend part of their PhD at the Donders Institute where Personalised Parkinson’s dataset has been acquired. This particular project will also offer the following specific training on signal processing of data obtained from wearable sensors and fMRI analysis.
Key Publications:
1. Cagnan, H. et al. Stimulating at the right time: phase-specific deep brain stimulation. Brain 140, 132–145 (2017).
2. Helmich, R. C., Janssen, M. J. R., Oyen, W. J. G., Bloem, B. R. & Toni, I. Pallidal dysfunction drives a cerebellothalamic circuit into Parkinson tremor. Annals of Neurology 69, 269–281 (2011).
3. Bloem, B. R. et al. The Personalized Parkinson Project: examining disease progression through broad biomarkers in early Parkinson’s disease. BMC Neurology 19, 160 (2019).
Funding: The successful applicant will be considered for a Clarendon fund or other University scholarship. Students with their own independent funding will also be considered.
Contact: Prof Hayriye Cagnan will be happy to discuss the project and PhD further. Please contact them by email on hayriye.cagnan@bndu.ox.ac.uk
Supervisor: Prof Peter Magill
Applications are invited from both Home students and International students to join a multidisciplinary team of scientists studying the neuronal cell and circuit substrates for purposeful movements in health and disease. Up to two MRC-funded studentships are available from the start of academic year 2021/22, for 3 years each, and both will be primarily supervised by Professor Peter Magill at the MRC BNDU.
Background: Release of the neurochemical dopamine in basal ganglia circuits helps us to correctly execute purposeful movements. In Parkinson’s disease and its animal models, the selective dysfunction and degeneration of midbrain dopaminergic neurons perturbs activity dynamics elsewhere in the basal ganglia to an extent that movement is profoundly impaired. It is important that we gain a detailed understanding of how dopaminergic neurons and other cell types in the basal ganglia fulfil their specialised roles to govern movement.
Project Description: The overall goal of these PhD studentship projects is to generate new mechanistic explanations of how the basal ganglia and their partner brain circuits work together to support normative movement in health as well as impaired movement in Parkinson’s disease. To achieve this, we will couple advanced analytical techniques with experimental interventions that capture and probe causal interactions between neurons with high spatiotemporal precision. The project will centre on the use of wild type and genetically-altered mice with intact or comprised midbrain dopaminergic systems, the readouts from which straddle multiple levels of function.
Training: The project will take place in the Medical Research Council Brain Network Dynamics Unit at the University of Oxford and students will benefit from the extensive generic and transdisciplinary skills training and career development opportunities available within the Unit. Focusing on the basal ganglia and partner brain circuits, studentship holders will harness cutting-edge methods in mice for identifying, monitoring, accessing and manipulating neurons in vivo. As an integral part of these projects, you will receive advanced training in several of the following research techniques; electrophysiology, anatomy, optogenetics, photometry, and mouse behavioural assays, together with rigorous data analysis. You will also be offered opportunities to work with sophisticated mouse models that capture genetic/proteinaceous burdens of relevance to Parkinson’s.
Key Publications:
Sharott A, Vinciati F, Nakamura KC, Magill PJ (2017) A population of indirect pathway striatal projection neurons is selectively entrained to Parkinsonian beta oscillations. J. Neurosci., 37(41):9977-9998.
Dodson PD, Dreyer JK, Jennings K, Syed EC, Wade-Martins R, Cragg SJ, Bolam JP, Magill PJ (2016) Representation of spontaneous movement by dopaminergic neurons is cell-type selective and disrupted in Parkinsonism. Proc. Natl. Acad. Sci. U.S.A., 113(15):E2180-2188.
Garas FN, Shah RS, Kormann E, Doig NM, Vinciati F, Nakamura KC, Dorst MC, Smith Y, Magill PJ, Sharott A (2016) Secretagogin expression delineates functionally-specialized populations of striatal parvalbumin-containing interneurons. eLife, 5:e16088.
Dodson PD, Larvin JT, Duffell JM, Garas FN, Doig NM, Kessaris N, Duguid IC, Bogacz R, Butt SJ, Magill PJ (2015) Distinct developmental origins manifest in the specialized encoding of movement by adult neurons of the external globus pallidus. Neuron, 86(2):501-513.
Abdi A, Mallet N, Mohamed FY, Sharott A, Dodson PD, Nakamura KC, Suri S, Avery SV, Larvin JT, Garas FN, Garas SN, Vinciati F, Morin S, Bezard E, Baufreton J, Magill PJ (2015) Prototypic and arkypallidal neurons in the dopamine-intact external globus pallidus. J. Neurosci., 35(17):6667-6688.
Funding: These Ph.D. (D.Phil.) studentships are funded by the Medical Research Council (MRC), a part of UKRI. All successful applicants are entitled to receive a tax-free stipend and, as a minimum, tuition fees paid at the Home level, regardless of whether they are Home or International students. Please see further details about MRC/UKRI studentships and updated guidance regarding Home and International eligibility. Successful applicants can also be considered for University scholarships.
Qualifications & Experience: Interested candidates should possess, or expect to receive, a 1st class or upper 2nd class degree (or equivalent) in a related scientific discipline, e.g. biological or physical sciences, medicine, computer science, engineering, mathematics. Previous experience in neuroscience research is highly desirable.
Contact: We strongly encourage candidates to contact the project supervisor before submitting an application. To find out more about these MRC-funded studentships, the research project, and the application process, please contact Professor Peter Magill by email on peter.magill@bndu.ox.ac.uk. Professor Magill is also happy to consider supervising PhD students funded by Departmental scholarships and other doctoral training programmes at Oxford, as well as PhD students with their own independent funding.
How to apply: To be considered for one of these MRC-funded studentships, please submit an application for admission to the D.Phil. in the Nuffield Department of Clinical Neurosciences (course code RD_CU1), following the guidance. On the application form, in the section headed ‘Departmental Studentship Applications’, please indicate that you are applying for a studentship and enter the reference code for a MRC BNDU studentship "21CLINNEURO01WEB".
The closing date for applications is 12.00 midday UK time on Friday 8th January 2021.
Interviews for short-listed applicants will be held between 25th and 28th January 2021.
Local field potential signal recorded from a patient with a deep brain stimulation electrode, showing a burst of beta-frequency oscillatory activity (top) and adaptive deep brain stimulation being triggered and reducing the length of another beta burst (bottom).
Applications & Funding
Applications & Funding
The MRC BNDU has its own MRC-funded Ph.D. (D.Phil.) studentships, which are typically awarded for three years in the first instance. Applicants to these MRC-funded studentships can also be considered for University scholarships. The Unit will also consider hosting PhD students funded by Departmental scholarships and other doctoral training programmes at Oxford. In addition, the Unit sometimes recruits for iCASE studentships in collaboration with industry. Information about University funding is available and an A-Z of Oxford scholarships. Students with their own independent funding will also be considered.
The MRC BNDU is one of the six divisions of the Nuffield Department of Clinical Neurosciences, and as such we benefit from the excellent student services and streamlined application process provided by the Department. The Nuffield Department of Clinical Neurosciences provides general information for prospective students, advice on entry requirements, and information about potential costs. There is also specific advice for prospective Home and International students.
Students considering applying for one or more of the PhD studentship projects available in the MRC BNDU should first contact the supervisor(s) named in the project description using the email provided in the project summary. You are encouraged to contact the supervisor(s) as soon as possible to find out more about the project and the application process. We look forward to hearing from you.
To be considered for a MRC-funded studentship following contact with the supervisor(s) of your proposed project, please submit an application following the online guidance for the D.Phil. in Clinical Neurosciences. On the application form, in the section headed ‘Departmental Studentship Applications’, you must indicate that you are applying for a studentship and enter the reference code for a MRC BNDU studentship.
In the special instance of an iCASE studentship, please contact the named supervisor(s) directly for advice on the application process.
Hippocampal theta oscillations (white) with associated faster rhythms (rainbow colours). From: Lopes-Dos-Santos V. et eal. Parsing Hippocampal Theta Oscillations by Nested Spectral Components during Spatial Exploration and Memory-Guided Behavior. 2018. Neuron, 100(4):940–952.