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Giraldez, T.

Publications and source records attributed to Giraldez, T..

8 recordsLinked to original sources

GRIN2B disease-associated mutations disrupt the function of BK channels and NMDA receptor signalling nanodomains

Large conductance calcium-activated potassium channels (BK channels) are unique in their ability to respond to two distinct physiological stimuli: intracellular Ca2+ and membrane depolarization. In neurons, these channels are activated through a coordinated response to both signals; however, for BK channels to respond to physiological voltage changes, elevated concentrations of intracellular Ca2+ (ranging from 1 to 10 M) are necessary. As a result, it is believed that BK channels are typically localized within nanodomains near Ca2+ sources (approximately 20-50 nm), such as N-methyl-D-aspartate receptors (NMDARs). Since the first evidence of NMDAR-BK channel coupling reported by Isaacson and Murphy in 2001 in the olfactory bulb, further studies have identified functional coupling between NMDARs and BK channels in other regions of the brain, emphasizing their importance in neuronal function. Mutations in the genes encoding NMDAR subunits have been directly linked to various neurodevelopmental disorders, including intellectual disability, epilepsy, and autism spectrum disorders. For instance, mutations such as V15M and V618G in the GRIN2B gene, which encodes the GluN2B subunit of NMDARs, are implicated in the pathogenesis of early infantile epileptic encephalopathy (EIEE27). Here, we explored the effects of these two GluN2B mutations on NMDAR-BK channel coupling, employing a combination of electrophysiological, biochemical, and imaging techniques. Taken together, our results demonstrate that mutation V618G specifically disrupts NMDAR-BK complex formation, impairing functional coupling, in spite of robust individual channel expression in the membrane. These results provide a potential mechanistic basis for EIEE27 pathophysiology and uncover new clues about NMDAR-BK complex formation.

physiology↗

State-dependent photocrosslinking unveiled the role of intersubunit interfaces on Ca2+ activating mechanisms of the BK channel

BK channels are high-conductance potassium channels that are activated by voltage and Ca2+. The pore-forming -subunits form homotetramers including a membrane-spanning domain and a cytosolic domain with a tandem of RCK-like domains (RCK1 and RCK2) per subunit. The eight RCKs compose high affinity Ca2+-binding sites that drive channel activation. Full-length Cryo-EM structures revealed intersubunit interactions between the RCK domains. Asparagine 449 (N449, human BK channel) is located at the RCK1 domain and coordinates Ca2+ in the RCK2 of the adjacent subunit. In addition, two Arginines (R786 and R790) of one RCK2 and Glutamate 955 of the adjacent subunit constitute an additional interaction interface. Functional studies on these residues showed that these two interfaces are crucial in Ca2+ sensitivity. To detect structural rearrangements induced by Ca2+ during channel activation, we took advantage of the photoactivatable unnatural amino acid p-benzoyl-L-phenylalanine (BzF). Functional channels were obtained with this amino acid inserted at 11 positions. N449 and R786 positions (N449BzF and R786BzF respectively). UV-induced photocrosslinking led to Ca2+ dependent and voltage-independent effects in both mutants. N449BzF showed a steady-state current reduction at saturating Ca2+ concentrations. Our data shows that this effect mainly relies on full occupancy of the RCK1 Ca2+ binding site, since mutation of this site abolished the effect. The R786BzF construct showed a substantial potentiation of the current in the absence of Ca2+. In this case, photocrosslinking seems to favor the activation of the channel by voltage. Overall, these results suggest mobile interfaces between RCK domains are key to BK channel activation.

biophysics↗

Structural bases for blockade and activation of BK channels by Ba2+ ions

Ba2+ increases open probability and blocks the permeation pathway of BK channels. Here we discuss Cryo-EM structures of BK channels with Ba2+. They reveal that Ba2+ occupies site S3 in the selectivity filter, leading to blockade. Ba2+ are also detected at all high affinity Ca2+ binding sites of the gating ring. However, the architectures of this region in these structures represent intermediate transitions between the extreme close and open configurations.

biophysics↗

Activation of SGK1.1 up-regulates the M-current in the presence of epilepsy mutations

In the central nervous system, the M-current plays a critical role in regulating subthreshold electrical excitability of neurons, determining their firing properties and responsiveness to synaptic input. The M-channel is mainly formed by subunits Kv7.2 and Kv7.3 that co-assemble to form a heterotetrametric channel. Mutations in Kv7.2 and Kv7.3 are associated with hyperexcitability phenotypes including benign familial neonatal epilepsy (BFNE) and neonatal epileptic encephalopathy (NEE). SGK1.1, the neuronal isoform of the serum and glucocorticoids-regulated kinase 1 (SGK1), increases M-current density in neurons, leading to reduced excitability and protection against seizures. Herein, using two-electrode voltage clamp on Xenopus laevis oocytes, we demonstrate that SGK1.1 selectively activates heteromeric Kv7 subunit combinations underlying the M-current. Importantly, activated SGK1.1 is able to up-regulate M-channel activity in the presence of two different epilepsy mutations found in Kv7.2 subunit, R207W and A306T. In addition, proximity ligation assays in the N2a cell line allowed us to address the effect of these mutations on Kv7-SGK1.1-Nedd4 molecular associations, a proposed pathway underlying M-channel up-regulation by SGK1.1

neuroscience↗

Synaptic plasticity of somatosensory neurons is determined by functional coupling between NMDA receptors and BK channels

Postsynaptic N-methyl-D-aspartate receptors (NMDARs) are crucial mediators of synaptic plasticity due to their ability to act as coincidence detectors of presynaptic and postsynaptic neuronal activity. However, NMDARs exist within the molecular context of a variety of postsynaptic signaling proteins, which can fine-tune their function. Here we describe a novel form of NMDAR suppression by large-conductance Ca2+- and voltage-gated K+ (BK) channels in the basal dendrites of a subset of barrel cortex layer 5 pyramidal neurons. We show that NMDAR activation increases intracellular Ca2+ in the vicinity of BK channels, thus activating K+ efflux and strong negative feedback inhibition. We further show that neurons exhibiting such NMDAR-BK coupling serve as high-pass filters for incoming synaptic inputs, precluding the induction of spike-timing-dependent plasticity. Together, these data suggest that NMDAR-localized BK channels regulate synaptic integration and provide input-specific synaptic diversity to a thalamocortical circuit.

neuroscience↗

Neuronal kinase SGK1.1 protects against brain damage after status epilepticus

Epilepsy is a neurological condition associated to significant brain damage produced by status epilepticus (SE) including neurodegeneration, gliosis and ectopic neurogenesis. Reduction of these processes constitutes a useful strategy to improve recovery and ameliorate negative outcomes after an initial insult. SGK1.1, the neuronal isoform of the serum and glucocorticoids-regulated kinase 1 (SGK1), has been shown to increase M-current density in neurons, leading to reduced excitability and protection against seizures. We now show that SGK1.1 activation potently reduces levels of neuronal death and gliosis after SE induced by kainate, even in the context of high seizure activity. This neuroprotective effect is not exclusively a secondary effect of M-current activation but is also directly linked to decreased apoptosis levels through regulation of Bim and Bcl-xL cellular levels. Our results demonstrate that this newly described antiapoptotic role of SGK1.1 activation acts synergistically with the regulation of cellular excitability, resulting in a significant reduction of SE-induced brain damage. The protective role of SGK1.1 occurs without altering basal neurogenesis in brain areas relevant to epileptogenesis. SIGNIFICANCE STATEMENTApproaches to control neuronal death and inflammation are of increasing interest in managing epilepsy, one of the most important idiopathic brain diseases. We have previously shown that activation of SGK1.1 reduces neuronal excitability by increasing M-current levels, significantly reducing seizure severity. We now describe a potent neuroprotective role of SGK1.1, which dramatically reduces neuronal death and gliosis after status epilepticus. This effect is partially dependent on M-current activation and includes an additional anti-apoptotic role of SGK1.1. Our data strongly support the relevance of this kinase as a potential target for epilepsy treatment.

neuroscience↗

SGK1.1 activation causes early seizure termination via effects on M-current

Early termination of status epilepticus affords protection against brain damage and associated pathologies. Regulation of Kv7.2/7.3 potassium channels, underlying the neuronal M-current, is key for seizure control. This conductance is maintained during initiation of action potentials, affecting neuronal excitability and thus inhibiting epileptic discharges. The M-current is upregulated by the neuronal isoform of the serum and glucocorticoid-regulated kinase SGK1 (SGK1.1). We tested whether SGK1.1 is an anticonvulsant factor using the kainic acid (KA) model of acute seizures in a transgenic mouse model with expression of a constitutively active form of the kinase. Our results demonstrate that SGK1.1 confers robust protection against seizures associated to lower mortality levels, independently of sex or genetic background. SGK1.1-dependent protection results in reduced number, shorter duration, and early termination of EEG seizures. At the cellular level, it is associated to increased M-current amplitude mediated by Nedd4-2 phosphorylation, leading to decreased excitability of hippocampal CA1 neurons without alteration of basal synaptic transmission. Altogether, our results reveal that SGK1.1-mediated M-current upregulation in the hippocampus is a key component of seizure resistance in the KA epileptic paradigm, suggesting that regulation of this anticonvulsant pathway may improve adverse outcomes to status epilepticus, constituting a potential target for antiepileptic drugs.

neuroscience↗

The BK channel gating ring is strongly coupled to the voltage sensor

The open probability of large conductance voltage- and calcium-dependent potassium (BK) channels is regulated allosterically by changes in the transmembrane voltage and intracellular concentration of divalent ions (Ca2+ and Mg2+). The divalent cation sensors reside within the gating ring formed by eight Regulator of Conductance of Potassium (RCK) domains, two from each of the four channel subunits. Overall, the gating ring contains 12 sites that can bind Ca2+ with different affinities. Using patch-clamp fluorometry, we have shown robust changes in FRET signals within the gating ring in response to divalent ions and voltage, which do not directly track open probability. Only the conformational changes triggered through the RCK1 binding site are voltage-dependent in presence of Ca2+. Because the gating ring is outside the electric field, it must gain voltage sensitivity from coupling to the voltage-dependent channel opening, the voltage sensor or both. Here we demonstrate that alterations of voltage sensor dynamics known to shift gating currents produce a cognate shift in the gating ring voltage dependence, whereas changing BK channels relative probability of opening had little effect. These results strongly suggest that the conformational changes of the RCK1 domain of the gating ring are tightly coupled to the voltage sensor function, and this interaction is central to the allosteric modulation of BK channels.

biophysics↗