region of projection cells and another adeno-associated virus vector (AAV2) in the region housing the cell bodies of these projection neurons (Kinoshita et al., 2012). The retrograde vector carried a gene for an enhanced tetanus neurotoxin (eTeNT) downstream of a tetracycline-responsive element (TRE). The AAV vector carried a reverse tetracycline transactivator (rtAV16). Hence, the eTeNT is conditionally expressed only in double-labeled neurons as it requires the transactivator. Moreover, only in the presence of doxycycline (a tetracycline), eTeNT will suppress synaptic transmission restricted to the double-infected projection neurons. In this first study, the technique revealed the critical role of propriospinal neurons in the control of hand dexterity in macaques. The same powerful double-infection technique was also instrumental to reveal the critical role of the connections between the superior colliculus and the ventrolateral pulvinar in blindsight after a V1 lesion (Kinoshita et al., 2019). Reversible inactivation of this pathway impaired visually guided saccades to the contralateral hemifield of blindsight monkeys.

In another recent study, the technique was used in combination with functional imaging in monkeys performing cognitive tasks (Vancraeyenest et al., 2020). Specifically, the authors reversibly inactivated the connections originating in the ventral tegmental area (VTA) and projecting to the nucleus accumbens (NAc) in monkeys. This predominantly dopaminergic pathway purportedly plays an important role in reinforcement-based learning. Contrary to this prediction, however, doxycycline-induced inactivation of the VTA-to-NAc pathway yielded no effect on reversal learning during an object reversal discrimination task. Instead, motivational behavior requiring high effort levels was severely diminished in another task. Moreover, reversible inactivation of the connections between the VTA and NAc resulted mainly in increased functional connectivity, measured with resting state fMRI, primarily between temporal and frontal regions. In the Vancraeyenest study, both HiRet (a vector pseudotyped with the fusion glycoprotein type B (FuG-B)) and NeuRet (with the fusion glycoprotein type C (FuG-C)) were used as retrograde gene transfer vectors. While both lentiviral vectors are highly efficient retrograde gene transfer vectors in primates, NeuRet possesses higher neuronal specificity than the HiRet vector (Kato et al., 2011). Recently, an even more efficient neuron-specific retrograde vector has been developed using another type of fusion glycoprotein (FuG-E) (Tanabe et al., 2017).

## 9.1. Discussion and outlook

The development of highly efficient retrograde transfer vectors in combination with conditional expression of genes to downregulate (e.g., the aforementioned neurotoxins or hyperpolarizing opsins) or enhance activity (e.g., in combination with depolarizing opsins) will offer fascinating new tools to dissect the contribution of specific neural pathways in perception and cognition (Inoue et al., 2015; Kinoshita et al., 2019, 2012; Nurminen et al., 2018; Tohyama et al., 2017). For translational purposes, these novel pathway-selective interventional approaches may have benefits over conventional region-specific methods, as they may mitigate unwanted side effects caused by the (in)activation of intersecting circuits at a target site that are not relevant for treatment of the disease.

Major challenges related to pathway-selective genetic based perturbation methods are highly similar to those described for optogenetics (see Section 6.3). It remains challenging to restrict diffusion of viral vectors exclusively to the intended targets, as is the case for all types of injections in the brain, including tract-tracing molecules, drugs, etc. A key control to reduce off-target effects is to combine the injection procedures with the most refined in-vivo imaging methods, for example using MRI while co-injecting MnCl₂ or GD-based contrast agents, or by combining injections with simultaneous electrophysiological recordings, or both.

The method can be used for studying the functionality of specific pathways at a mesoscale level, such as investigating functional inter-

actions between blobs in area V1 and different stripe compartments in area V2, provided the dual injections can be confined to these specific functional subcompartments. This will most likely require the use of optical imaging or focal ultrasound imaging methods to identify the exact targets (i.e., in the example above, those would be the blobs in V1 and a specific stripe compartment in V2). At macroscale levels, the technology can be used to investigate functional interactions across areas, such as interactions between the FEF and area V4. Both the meso- and macroscale applications can be rendered cell-type specific when additional promoters in the vector are inserted to target these cell types (e.g., TH promoter to target dopaminergic neurons). An obvious limitation is that viral vectors which are cell-type selective and also efficient to use, are currently still very limited in availability and not used much for NHP research.

Theoretically, the reversible pathway-selective perturbation methods described above would be applicable for human use, since viral vector-based treatments are currently already implemented in a variety of clinical applications. Obvious safety protocols should be followed, as is the case with all clinical viral vector methods. The advantage of the dual injection technique compared to, for example, optogenetics is that after injection of the viral vectors, no additional invasive procedures are required to 'activate' or 'inactivate' the targeted pathway. Because of the presence of a reverse transactivator in combination with the tetracycline-responsive element, only a tetracycline needs to be administered (e.g. orally) to activate the downstream gene (in the example above, this would be eTeNT). It is conceivable to insert different genes downstream relative to the tetracycline-responsive element, whereby a specific pathway can be reversibly boosted instead of silenced, as long as the tetracycline is given to the patient. Such pathway-selective clinical interventions would offer the possibility to focally target (inactivate or activate) a specific pathway within a complex system affected by a specific disease and then assess improvement in symptoms. As an example, this approach could be used to evaluate an under- or over-performing dopaminergic reward system in the case of depression or addiction, respectively. The approach could be more powerful and induce fewer side effects because it would not impact off-target pathways that may be highly relevant to support normal behavior unaffected by the disease.

## 10. General discussion

Brain perturbation methods are vital for advancing understanding about the causal mechanisms of distributed brain functions. Employing brain perturbation techniques in NHPs has been and will continue to be instrumental for scientific and biomedical advances. Combined with neuroimaging, these techniques and the insights gained with them can become even more powerful, allowing brain-wide visualisation of the perturbation impact on brain systems and interconnectivity. Neuroimaging also allows comparison of effects across methodologically distinct techniques. Such comparisons enhance insights into the mode of action of the perturbation techniques, which is vital for understanding how they can optimally translate to and benefit humans. The interplay between neuroimaging and brain perturbation not only improves our understanding of the brain perturbation techniques, but it can also provide novel insights into the neural underpinnings of the neuroimaging signals themselves (e.g., the MRI-BOLD signal).

This review could not cover every brain perturbation technique that has been used in combination with neuroimaging. It focuses on several of the more commonly used techniques that are regularly combined with neuroimaging, and highlights some new approaches with exciting potential. Other brain perturbation approaches like cooling (Khachaturian et al., 2008), Transcranial Magnetic Stimulation (TMS) (Balan et al., 2017), or transcranial current stimulation (tDCS/tACS) (see Liu et al., 2018), and references therein) are less commonly used in NHPs, even though they are regularly combined with neuroimaging in humans (e.g., TMS-fMRI). These methods may be less commonly studied in NHPs for both practical reasons and because of the greater specificity
of other approaches. Nonetheless, even for these or other approaches, work with NHPs may be required to understand the underlying neural mechanisms and the combination with neuroimaging could be an important component (Balan et al., 2017).

This paper offers a snapshot of where the field stands in terms of NHP brain perturbation and neuroimaging advances. However, because many developments are ongoing, we envision a dynamic resource on brain perturbation and neuroimaging that evolves along with advances in the field as a complementary resource to this paper. The PRIME-RE platform (Messinger et al., 2021), that was recently launched by members of the PRIME-DE community as an online research exchange platform for all things related to NHP neuroimaging. It offers an excellent outlet for such dynamically evolving content and will encourage data sharing of legacy and new data, as well as offering primers to support laboratories to use and develop the techniques further. Maintained by the community, researchers can upload technique primers, tips, tricks, risks, solutions and warnings for all relevant methodologies. This ongoing effort will help the scientific community to keep up to date with recent developments and to support researchers aiming to implement the techniques in their own labs. PRIME-RE also maintains collections of links to open NHP neuroimaging datasets and could facilitate the distribution and accessibility of valuable perturbation datasets, that might for instance be hosted on PRIME-DE (Milham et al., 2018).

## 10.1. Anesthesia as a perturbation method

The implications of performing brain perturbation and neuroimaging studies in either anesthetized or awake animals were briefly mentioned here and more extensively detailed for NHP studies elsewhere (Basso et al., 2021). However, anesthesia has non-uniform effects on the brain, and different anesthetic agents impact different systems in the brain (e.g., pain neuraxis, consciousness). For instance, anesthesia suppresses the feedback of neural processing (Suzuki and Larkum, 2020), and some brain regions, such as the precuneus, posterior cingulate gyrus, prefrontal dorsolateral cortex and thalamus, are more strongly deactivated by various forms of anesthesia than other regions (Franks, 2008). The non-uniform effects of anesthesia have been exploited in studies on the neural mechanisms of auditory sequence violations (Bekinschtein et al., 2009). For instance, neural responses to oddball sounds (local violations) primarily engage earlier auditory processing stages, while relationships that require between sequences integration of information (global violations) engage broader brain circuits, including the prefrontal, parietal and cingulate cortices (Uhrig et al., 2014). The local violation effect in the auditory cortex disappears under propofol anesthesia (a GABA-ergic agonist) and, interestingly, shifts spatially under ketamine anesthesia (an NMDA antagonist) (Uhrig et al., 2016). A progressive disorganization of the global effect in the prefrontal, parietal and cingulate cortex was observed with increasing levels of propofol anesthesia, while ketamine completely suppressed these same areas. Anesthesia has also been shown to massively reconfigure dynamic resting-state networks (Barttfeld et al., 2015; Uhrig et al., 2018) despite a preservation of stationary resting-state networks (Vincent et al., 2007). Although anesthesia is not usually thought of as a brain perturbation approach, using neuroimaging to understand anesthetic agent mechanisms and visualize the impact on the brain is important scientifically as well as for clinical and other translational reasons.

## 10.2. The perturbed brain as a functional model

One important limitation of perturbation studies is that adding or removing activity may force the brain into a state that might not occur naturally. On the other hand, brain impact or disease is by definition not normal. Thus, one could question to what extent the resulting effects are representative of the brain's normal mode of operation. In a sense, this

question is not categorically different from questioning whether artificial stimuli employed in a laboratory environment can teach us anything about how the brain operates in natural environments. In this case, the input is much more complex, comprehensive and likely to simultaneously engage many processing networks, leading to a broad range of complex neural interactions (Rust and Movshon, 2005). Any precisely targeted brain perturbation is likely to modulate only a fraction of the networks typically involved in 'natural' perception, cognition, and behavior. As a consequence, drawing causal conclusions at the systems level after perturbation of a single node in a complex network should be done with caution, especially when asserting necessity or sufficiency. Future NHP studies could also perturb multiple nodes in the network to more rigorously and thoroughly assess causal relationships (Qiao et al., 2020), a unique approach that is not possible with humans. However, a reductionist approach with either sensory stimulation or brain perturbation can be an advantage for teasing apart the mechanisms of a complex system that is the brain; so long as we remain aware of the fact that we are introducing a potentially unnatural brain perturbation. On the other hand, artificial stimuli can be designed to emulate natural conditions and likewise, with increasing understanding of their underlying mechanisms, the propagation of brain perturbation effects can become increasingly similar to natural signal propagation. In this regard, the spatio-temporal scope of the intended perturbation approach (Fig. 1) will allow selecting the most useful tool to use for the intended purpose. Careful comparison of brain dynamics in unperturbed and perturbed situations furthermore offers the advantage of studying the system in a more natural state or during perturbation, as natural or unnatural as that might be. After all, brain damage in patients is akin to a brain perturbation technique that disrupts normal functioning, with the experimental approach in NHPs as a model system having the advantage of being able to perturb the brain in a highly controlled way. Thus, although this paper underscores the value of brain perturbation approaches, the benefits of comparing results to descriptive studies without perturbation should not be dismissed.

## 10.3. Causality and comparison across perturbation techniques

We have emphasized that a key benefit of brain perturbation approaches is that they allow inferences on cause and effect. The various techniques can provide different, partially non-overlapping ranges of spatial and temporal resolution, and cell type- or pathway-specificity (Fig. 1), but there are several conditions that need to be met to obtain true causal information (Clark et al., 2011). As discussed in the technique-specific sections above, the confidence with which causality can be inferred depends on the level of understanding of the mechanism of action of a particular technique as well as its spatio-temporal scope. Even for techniques for which there is a substantial evidence base, this type of knowledge is still incomplete. A second limitation of any perturbation study is that even if a causal relationship can be asserted with relative confidence, it does not necessarily mean that this specific causal relation is sufficient or necessary to support a brain function or behavior (see also Section 10.2). A thorough comparison across different studies and perturbation methods, as well as approaches in which multiple nodes are perturbed simultaneously, may also shed light on this issue. Thus, it is crucial to continue to increase our understanding of the specificity and modes of action of individual perturbation approaches, as well as the strengths and limitations of different neuroimaging methods. Using neuroimaging to both record and compare the impact of different brain perturbation techniques will continue to shed light on the different modes of action and the types of causal inferences that can be made.

## 10.4. Suppressed or negative imaging signals

Functional imaging studies often tend to focus on positive brain signals, but neural processes can manifest as either increases or decreases of the neuroimaging signal. This is not unique to fMRI imaging but also
applies to other neuroimaging modalities such as PET. Although base-
line levels of brain activity can fluctuate and be difficult to define, both
positive and negative effects can be crucial for understanding the neu-
ral mechanisms and mode of action of a brain perturbation approach.
Yet, to rectify what may be a bias towards reporting positive responses,
it is important to consider the neural basis of negative imaging signals.
Intuitively, they could be related to neuronal suppression, but in some
cases they could also reflect an entirely different process, which may be
specific to the mode of action of a particular technique.

Sensory stimulation leads to reliable positive and negative fMRI
BOLD responses (PBRs and NBRs) during topographical mapping
(Allison et al., 2000; Fracasso et al., 2018; Harel et al., 2002; Klink et al.,
2020). NBRs can occur next to PBRs in the visual cortex and have
been shown to correlate with electrophysiologically-measured neuronal
inhibition (Huang et al., 1996; Shmuel et al., 2006, 2002). NBRs
are also found with frequency mapping of the auditory cortex, adja-
cent to areas of PBRs (Ortiz-Rios et al., 2017; Petkov et al., 2006;
Tanji et al., 2010). Local inhibitory lateral connections (Angelucci and
Bullier, 2003) as well as distal excitatory and inhibitory interactions
might play a role in NBR effects through a push/pull mechanism
(Angelucci and Bressloff, 2006). Although positive BOLD or MION-based
fMRI responses tend to predominate during cortical electrical micros-
timulation, suppression of fMRI responses appears to be common in the
few available NHP studies with subcortical stimulation. The presence of
NBRs may depend on stimulation frequency, animal state (e.g., awake
or anesthetized) and the intrinsic properties of the stimulated region
(Logothetis et al., 2010; Murayama et al., 2011; Murris et al., 2020;
Sultan et al., 2011).

Some negative imaging signal effects are not due to neuronal re-
sponse suppression, and neuronal response suppression can even re-
sult in a positive fMRI signal if it increases neurovascular demands
(e.g., greater responses by inhibitory neurons that require a vascular
response). The nature of negative fMRI signals can furthermore dif-
fer substantially between BOLD-based fMRI and MION contrast-agent-
based approaches (Leite et al., 2002; Mandeville, 2012; Vanduffel et al.,
2001). MION is more commonly used with macaque 3T imaging for its
superior contrast to noise properties and because it is a cleaner measure
of cerebral blood volume (CBV) than BOLD-fMRI. Yet, hemodynamic
differences between BOLD- and MION-based fMRI could be exploited
to further study the nature of neurovascular coupling. For instance,
Yu et al. (2016) found that MION-CBV resulted in earlier onset responses
from arterioles and BOLD-fMRI resulted in later onset responses from
venules. With new opsin development allowing the manipulation of neu-
ronal activity using inhibition (Li et al., 2019), and the perturbation of
transsynaptic circuits (Tervo et al., 2016), optogenetics combined with
functional imaging could be used to clarify the neural bases of positive
and negative fMRI effects.

There may also be technique-specific reasons for negative imaging
signals. For instance, in optogenetics studies, high-power photostimu-
lation can generate artifactual negative responses that are related to
the high absorption of light by the local neural tissue, thought to re-
sult from intravoxel temperature gradients changes and in a decrease of
the T2* signal (Albers et al., 2019). This particular effect can partially
be resolved by using red-shifted opsin s which allow the use of longer
wavelength light for stimulation (red instead of blue), which exhibits
less tissue absorption and moderate scattering. Widespread distal nega-
tive responses have been reported in rodent studies of optogenetically
stimulated cortex (Chan et al., 2017), but the neuronal bases for such
effects require further study and a corresponding primate optogenetics
study.

Neuronal recordings and high-resolution imaging could help to illu-
minate the bases for both positive and negative imaging effects. With
the increasingly regular use of higher magnetic field strength MRI scan-
ners in humans, it is now becoming possible to functionally differ-
entiate signals from different sets of cortical layers in both humans
(Huber et al., 2020, 2014) and animals (Chen et al., 2019a,b), and it

is evident that MION-CBV and BOLD-fMRI responses can pick up dif-
ferent signals across the cortical layers (Goense et al., 2012; Jin and
Kim, 2008; Smirnakis et al., 2007). Although highly ambitious, future
high field perturbation neuroimaging studies combined with laminar
neurophysiological recordings could fundamentally improve our under-
standing of brain systems and neural mechanisms of brain perturbation
effects.

10.5. Translational potential rooted in a solid fundamental science
foundation

Perturbation techniques in NHPs have substantial translational po-
tential. This is discussed at length for the different techniques in the Dis-
cussion and Outlook sections above, but we will briefly summarize and
present an overview here. The obtained knowledge about causal brain
mechanisms and the development of safe and effective methodologies
could all inform and improve treatment options in humans. Some per-
turbation approaches complement methodologies that are already being
used in humans and provide vital information that would be very dif-
ficult or impossible to obtain in humans. Other approaches may soon
have greater use in humans.

The development of DBS in humans as a treatment option for de-
bilitating disorders substantially benefited from electrical stimulation
studies in NHPs and will continue to do so as further information on
mode of action and brain impact remains needed. Now that DBS is
also being used to treat neuropsychiatric disorders, or to enhance phar-
macological treatment, the combination of neuroimaging and electri-
cal/pharmacological perturbation in controlled NHP studies is an in-
dispensable part of the translational pipeline. Its study in NHPs could
maximize clinical translational potential and benefits, give rise to suc-
cessful translational pipelines from rodents to primates to humans, and
incidentally, in the other direction as well.

Other brain perturbation approaches (e.g., permanent or reversible
lesions, pharmacological and genetic manipulations) also have well-
established links to human clinical research, while some of the newer
techniques (e.g., ultrasound, infrared stimulation) are at different stages
of translation to humans. For example, while FUS is already being used
in humans, this application firmly depends on information that is being
obtained with rodents and NHPs concerning its mode of action, impact
on neural systems, and optimal parameters for effective and safe use
(including blood-brain barrier manipulations).

Although translation to humans is an important goal of biomedical
science, advancing scientific knowledge is an equally, if not an even
more important objective. Moreover, clinical translation may simply
not be possible, fail, or be unsafe for humans without a firm bedrock
of knowledge provided by fundamental animal research. Comparative
neuroimaging studies are an example of fundamental science that fo-
cuses primarily on advancing the kind of scientific knowledge that is cru-
cial for understanding the evolution of neural systems and identifying
evolutionary conservation and divergence across species. Such under-
standing is vital to take into account when translating scientific results
and knowledge between animal model systems and humans (Mars et al.,
2018; Van Essen and Dierker, 2007; Xu et al., 2020a,b).

Another important limitation of any translational or preclinical
study, including those involving brain perturbations, is the biological
variability in both the (clinical) population and the sample of subjects or
animals enrolled in the study. For NHP studies that are often performed
with relatively small sample sizes for ethical or practical reasons, this
becomes particularly challenging when results are projected to a hetero-
geneous clinical population of a poorly understood disorder. One partial
solution to this challenge is to design studies that address basic functions
that can be related to symptoms, but are only partial disease or disorder
models. This approach is built around the assumption that basic func-
tions and their biological substrates are less variable than dysfunctions
or disorders, which are often diagnosed based on heterogeneous collec-
tions of symptoms. An improved understanding of a functioning system
can thus provide the bedrock for designing or improving treatments for dysfunctional neural systems. This approach is also a core component of the Research Domain Criteria (RDoC) framework suggested for the study of mental disorders (Insel et al., 2010). The sharing and combining of datasets and methods is another way to increase sample sizes and facilitate meta-analyses. The PRIME-DE (Milham et al., 2018; The PRIMatE Data Exchange (PRIME-DE) Global Collaboration Workshop and Consortium et al., 2020) and PRIME-RE (Messinger et al., 2021) initiatives are specifically designed to facilitate these efforts for NHP neuroimaging research.

Finally, rather than assuming that any primate brain will be a good model for the human brain, basic science studies, including cross-species comparative neuroimaging, will continue to be needed to assess this assumption by explicitly testing for correspondence or divergence across species (Orban et al., 2004; Rocchi et al., 2021; Tootell et al., 2003; Vanduffel et al., 2014). For example, cross-species alignment of cortical receptor and gene expression revealed highly similar patterns of serotonin 5HT<sub>1A</sub> receptors in humans and macaque monkeys, and weaker correspondence with rats (Froudist-Walsh et al., 2021). This is promising for NHP models of depression, as conventional antidepressants increase 5HT<sub>1A</sub> receptor signalling (Carhart-Harris and Nutt, 2017). Moreover, growing evidence suggests that the neural substrate of some cognitive functions is mostly symmetrical across the two brain hemispheres in NHPs, whereas substantial hemispheric lateralization exists for similar functions in the human brain (Balezeau et al., 2020; Hutchison et al., 2012a). Such differences between NHPs and humans have been identified in hemispheric lateralization of the fronto-parietal networks involved in spatial attention (Kagan et al., 2010; Mantini et al., 2013; Patel et al., 2015; Wey et al., 2014). The identified differences between humans and NHPs should be better understood, and considered alongside the evidence for shared evolutionarily conserved principles of primate brain organization. As we have noted here, NHP research often converges with and informs work in human patients. For instance, the compensatory, beneficial involvement of the intact hemisphere after unilateral lesions in NHPs (Sections 2 and 3) is in line with patient studies that demonstrate the recruitment of the opposite hemisphere during recovery after unilateral brain damage (Bartolomeo and Thiebaut de Schotten, 2016; Umarova et al., 2016). Likewise, the role of areas along the superior temporal sulcus in spatial orienting and attention that emerges from NHP studies is paralleled by a recent emphasis on temporal and temporal-parietal damage in neglect patients (Karnath and Rorden, 2012).

The exquisite technological approaches, along with the corresponding expansion of knowledge they have generated, illustrate that the field of combined brain perturbation and neuroimaging in non-human primates carries tremendous promise for science and medicine. It has, and will continue to have, a distinct, important, and lasting influence on both fundamental and translational neuroscience.

## No data statement

This paper does not contain original data or analysis.

## Declaration of Competing Interest

The authors declare no competing financial interests.

## Acknowledgements

This work was supported by NIH-MH112142 (V.F. and E.K.); California National Primate Research Center Base Grant (P51-OD011107) (A.F.); Fondation Bettencourt Schueller, France and Human Brain Project (B.J., L.U., J.T.); the Intramural Research Program of the NIMH (ZIA MH002918) (A.M.); Wellcome Trust (WT 110157/Z/15/Z) (A.S.M.); European Research Council (ERC StG, OptoVision, 637638) (M.C.S and M.O.R.); National Key R&D Program

of China 2018YFA0701400 (A.W.R.); Chinese NSF (81430010 and 31627802) (A.W.R.); Zhejiang province 2020C03004 (A.W.R.); Chinese National R&D Program Grant 2015AA020515 (A.W.R.); IDEXYLYON "IMPULSION 2020 grant (IDEX/IMP/2020/14) (J.S.); Howard Hughes Medical Institute (D.T.); KU Leuven grant C14/17/109 (W.V.); the National Eye Institute Intramural Research Program (ZIA EY000511) at the National Institutes of Health, U.S.A. (W.V.); Fonds Wetenschappelijk Onderzoek-Vlaanderen (FWO) G0D5817N, G0B8617N, G0C1920N, G0E0520N, VS02219N (W.V.); the European Union's Horizon 2020 Framework Programme for Research and Innovation under Grant Agreement No 945539 (Human Brain Project SGA3) (W.V.); Hermann and Lilly Schilling Foundation; German Research Foundation (DFG) grants WI 4046/1-1 and Research Unit GA1475-B4, KA 3726/2-1, Primate Platform of DFG Center for Nanoscale Microscopy & Molecular Physiology of the Brain (CNMPB) (M.W. and I.K.); Wellcome Trust (WT092606AIA) (C.I.P.); European Research Council Horizon 2020 (ERC CoG, MECHIDENT 724198) (C.I.P.); the Bettencourt Schueller Foundation (Ultrabrain project) (J.-F. A.); the French National Agency for Research (ANR-10-EQPX-15) (J.-F. A.); and the National Institutes of Health (with Matthew Howard III: R01-DC04290) (C.I.P.);

All research discussed in this work was performed in accordance with institutional and nationally approved oversight, such as the NIH Guide for Care and Use of Laboratory Animals, the U.K. Animals (Scientific Procedures) Act, 1986, and European Directive 2010/63/EU.

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