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One-carbon/folate (1C) metabolism supplies methyl groups required for DNA and histone methylation, and is involved in the maintenance of self-renewal in stem cells. Dihydrofolate reductase (DHFR), a key enzyme in 1C metabolism, is highly expressed in human and mouse neural progenitors at the early stages of neocortical development. Here, we have investigated the role of DHFR in the developing neocortex and report that reducing its activity in human neural organoids and mouse embryonic neocortex accelerates indirect neurogenesis, thereby affecting neuronal composition of the neocortex. Furthermore, we show that decreasing DHFR activity in neural progenitors leads to a reduction in one-carbon/folate metabolites and correlates with modifications of H3K4me3 levels. Our findings reveal an unanticipated role for DHFR in controlling specific steps of neocortex development and indicate that variations in 1C metabolic cues impact cell fate transitions.
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Neocórtex , Neurogénesis , Tetrahidrofolato Deshidrogenasa , Animales , Humanos , Ratones , Carbono , Ácido Fólico , Neurogénesis/genética , Tetrahidrofolato Deshidrogenasa/genéticaRESUMEN
Pseudocapacitors bridge the performance gap between batteries and electric double-layer capacitors by storing energy via a combination of fast surface/near-surface Faradaic redox processes and electrical double-layer capacitance. Organic semiconductors are an emerging class of pseudocapacitive materials that benefit from facile synthetic tunability and mixed ionic-electronic conduction. Reported examples are mostly limited to p-type (electron-donating) conjugated polymers, while n-type (electron-accepting) examples remain comparatively underexplored. This work introduces a new cross-linked n-type conjugated polymer, spiro-NDI-N, strategically designed with polar tertiary amine side chains. This molecular design aims to synergistically increase the electroactive surface area and boost ion transport for efficient ionic-electronic coupling. Spiro-NDI-N demonstrates excellent pseudocapacitive energy storage performance in pH-neutral aqueous electrolytes, with specific capacitance values of up to 532 F g-1 at 5 A g-1 and stable cycling over 5000 cycles. Moreover, it maintains a rate capability of 307 F g-1 at 350 A g-1. The superior pseudocapacitive performance of spiro-NDI-N, compared to strategically designed structural analogues lacking either the cross-linked backbone or polar side chains, validates the essential role of its molecular design elements. More broadly, the design and performance of spiro-NDI-N provide a novel strategy for developing high-performance organic pseudocapacitors.
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Electrochemical detection of metabolites is essential for early diagnosis and continuous monitoring of a variety of health conditions. This review focuses on organic electronic material-based metabolite sensors and highlights their potential to tackle critical challenges associated with metabolite detection. We provide an overview of the distinct classes of organic electronic materials and biorecognition units used in metabolite sensors, explain the different detection strategies developed to date, and identify the advantages and drawbacks of each technology. We then benchmark state-of-the-art organic electronic metabolite sensors by categorizing them based on their application area (in vitro, body-interfaced, in vivo, and cell-interfaced). Finally, we share our perspective on using organic bioelectronic materials for metabolite sensing and address the current challenges for the devices and progress to come.
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Técnicas Biosensibles , Electrónica , Compuestos OrgánicosRESUMEN
The organic electrochemical transistor (OECT) is one of the most versatile devices within the bioelectronics toolbox, with its compatibility with aqueous media and the ability to transduce and amplify ionic and biological signals into an electronic output. The OECT operation relies on the mixed (ionic and electronic charge) conduction properties of the material in its channel. With the increased popularity of OECTs in bioelectronics applications and to benchmark mixed conduction properties of channel materials, the characterization methods have broadened somewhat heterogeneously. We intend this review to be a guide for the characterization methods of the OECT and the channel materials used. Our review is composed of two main sections. First, we review techniques to fabricate the OECT, introduce different form factors and configurations, and describe the device operation principle. We then discuss the OECT performance figures of merit and detail the experimental procedures to obtain these characteristics. In the second section, we shed light on the characterization of mixed transport properties of channel materials and describe how to assess films' interactions with aqueous electrolytes. In particular, we introduce experimental methods to monitor ion motion and diffusion, charge carrier mobility, and water uptake in the films. We also discuss a few theoretical models describing ion-polymer interactions. We hope that the guidelines we bring together in this review will help researchers perform a more comprehensive and consistent comparison of new materials and device designs, and they will be used to identify advances and opportunities to improve the device performance, progressing the field of organic bioelectronics.
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A promising class of materials for applications that rely on electron transfer for signal generation are the n-type semiconducting polymers. Here we demonstrate the integration of an n-type conjugated polymer with a redox enzyme for the autonomous detection of glucose and power generation from bodily fluids. The reversible, mediator-free, miniaturized glucose sensor is an enzyme-coupled organic electrochemical transistor with a detection range of six orders of magnitude. This n-type polymer is also used as an anode and paired with a polymeric cathode in an enzymatic fuel cell to convert the chemical energy of glucose and oxygen into electrical power. The all-polymer biofuel cell shows a performance that scales with the glucose content in the solution and a stability that exceeds 30 days. Moreover, at physiologically relevant glucose concentrations and from fluids such as human saliva, it generates enough power to operate an organic electrochemical transistor, thus contributes to the technological advancement of self-powered micrometre-scale sensors and actuators that run on metabolites produced in the body.
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Fuentes de Energía Bioeléctrica , Técnicas Biosensibles , Técnicas Electroquímicas , Glucosa/metabolismo , Saliva/metabolismo , HumanosRESUMEN
Donor-acceptor (D-A) polymers are promising materials for organic electrochemical transistors (OECTs), as they minimize detrimental faradaic side-reactions during OECT operation, yet their steady-state OECT performance still lags far behind their all-donor counterparts. We report three D-A polymers based on the diketopyrrolopyrrole unit that afford OECT performances similar to those of all-donor polymers, hence representing a significant improvement to the previously developed D-A copolymers. In addition to improved OECT performance, DFT simulations of the polymers and their respective hole polarons also reveal a positive correlation between hole polaron delocalization and steady-state OECT performance, providing new insights into the design of OECT materials. Importantly, we demonstrate how polaron delocalization can be tuned directly at the molecular level by selection of the building blocks comprising the polymers' conjugated backbone, thus paving the way for the development of even higher performing OECT polymers.
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Generation of glial cell diversity in the developing spinal cord is known to depend on spatio-temporal patterning programs. In particular, expression of the transcription factor Olig2 in neural progenitors of the pMN domain is recognized as critical to their fate choice decision to form oligodendrocyte precursor cells (OPCs) instead of astrocyte precursors (APs). However, generating some confusion, lineage-tracing studies of Olig2 progenitors in the spinal cord provided evidence that these progenitors also generate some astrocytes. Here, we addressed the role of the heparan sulfate-editing enzyme Sulf2 in the control of gliogenesis and found an unanticipated function for this enzyme. At initiation of gliogenesis in mouse, Sulf2 is expressed in ventral neural progenitors of the embryonic spinal cord, including in Olig2-expressing cells of the pMN domain. We found that sulf2 deletion, while not affecting OPC production, impairs generation of a previously unknown Olig2-expressing pMN-derived cell subtype that, in contrast to OPCs, does not upregulate Sox10, PDGFRα or Olig1. Instead, these cells activate expression of AP identity genes, including aldh1L1 and fgfr3 and, of note, retain Olig2 expression as they populate the spinal parenchyma at embryonic stages but also as they differentiate into mature astrocytes at postnatal stages. Thus, our study, by revealing the existence of Olig2-expressing APs that segregate early from pMN cells under the influence of Sulf2, supports the existence of a common source of APs and OPCs in the ventral spinal cord and highlights divergent regulatory mechanism for the development of pMN-derived OPCs and APs.
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Astrocitos/enzimología , Factor de Transcripción 2 de los Oligodendrocitos/metabolismo , Médula Espinal/enzimología , Sulfatasas/metabolismo , Animales , Astrocitos/citología , Sustancia Gris/citología , Sustancia Gris/enzimología , Sustancia Gris/crecimiento & desarrollo , Ratones Transgénicos , Células-Madre Neurales/citología , Células-Madre Neurales/enzimología , Neurogénesis/fisiología , Receptor Tipo 3 de Factor de Crecimiento de Fibroblastos/metabolismo , Factores de Transcripción SOXE/metabolismo , Médula Espinal/citología , Médula Espinal/crecimiento & desarrollo , Sulfatasas/genéticaRESUMEN
A subclass of organic semiconductors known as conjugated polyelectrolytes (CPEs) is characterized by a conjugated backbone with ionic pendant groups. The water solubility of CPEs typically hinders applications of thin films in aqueous media. Herein, it is reported that films of an anionic CPE, namely CPE-K, drop cast from water produces single-component solid-state pseudocapacitive electrodes that are insoluble in aqueous electrolyte. That X-ray diffraction experiments reveal a more structurally ordered film, relative to the as-obtained powder from chemical synthesis, and dynamic light scattering measurements show an increase in aggregate particle size with increasing [KCl] indicate that CPE-K films are insoluble because of tight interchain contacts and electrostatic screening by the electrolyte. CPE-K film electrodes can maintain 85% of their original capacitance (84 F g-1 ) at 500 A g-1 and exhibit excellent cycling stability, where a capacitance retention of 93% after 100 000 cycles at a current density of 35 A g-1 . These findings demonstrate that it is possible to use initially water soluble ionic-organic materials in aqueous electrolytes, by increasing the electrolyte concentration. This strategy can be applied to the application of conjugated polyelectrolytes in batteries, organic electrochemical transistors, and electrochemical sensors, where fast electron and ion transport are required.
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The ability to amplify, translate, and process small ionic potential fluctuations of neural processes directly at the recording site is essential to improve the performance of neural implants. Organic front-end analog electronics are ideal for this application, allowing for minimally invasive amplifiers owing to their tissue-like mechanical properties. Here, we demonstrate fully organic complementary circuits by pairing depletion- and enhancement-mode p- and n-type organic electrochemical transistors (OECTs). With precise geometry tuning and a vertical device architecture, we achieve overlapping output characteristics and integrate them into amplifiers with single neuronal dimensions (20 micrometers). Amplifiers with combined p- and n-OECTs result in voltage-to-voltage amplification with a gain of >30 decibels. We also leverage depletion and enhancement-mode p-OECTs with matching characteristics to demonstrate a differential recording capability with high common mode rejection rate (>60 decibels). Integrating OECT-based front-end amplifiers into a flexible shank form factor enables single-neuron recording in the mouse cortex with on-site filtering and amplification.
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Due to their effective ionic-to-electronic signal conversion and mechanical flexibility, organic neural implants hold considerable promise for biocompatible neural interfaces. Current approaches are, however, primarily limited to passive electrodes due to a lack of circuit components to realize complex active circuits at the front-end. Here, we introduce a p-n organic electrochemical diode using complementary p- and n-type conducting polymer films embedded in a 15-µm -diameter vertical stack. Leveraging the efficient motion of encapsulated cations inside this polymer stack and the opposite doping mechanisms of the constituent polymers, we demonstrate high current rectification ratios ([Formula: see text]) and fast switching speeds (230 µs). We integrate p-n organic electrochemical diodes with organic electrochemical transistors in the front-end pixel of a recording array. This configuration facilitates the access of organic electrochemical transistor output currents within a large network operating in the same electrolyte, while minimizing crosstalk from neighboring elements due to minimized reverse-biased leakage. Furthermore, we use these devices to fabricate time-division-multiplexed amplifier arrays. Lastly, we show that, when fabricated in a shank format, this technology enables the multiplexing of amplified local field potentials directly in the active recording pixel (26-µm diameter) in a minimally invasive form factor with shank cross-sectional dimensions of only 50×8 [Formula: see text].
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Introducing ethylene glycol (EG) side chains to a conjugated polymer backbone is a well-established synthetic strategy for designing organic mixed ion-electron conductors (OMIECs). However, the impact that film swelling has on mixed conduction properties has yet to be scoped, particularly for electron-transporting (n-type) OMIECs. Here, the authors investigate the effect of the length of branched EG chains on mixed charge transport of n-type OMIECs based on a naphthalene-1,4,5,8-tetracarboxylic-diimide-bithiophene backbone. Atomic force microscopy (AFM), grazing-incidence wide-angle X-ray scattering (GIWAXS), and scanning tunneling microscopy (STM) are used to establish the similarities between the common-backbone films in dry conditions. Electrochemical quartz crystal microbalance with dissipation monitoring (EQCM-D) and in situ GIWAXS measurements reveal stark changes in film swelling properties and microstructure during electrochemical doping, depending on the side chain length. It is found that even in the loss of the crystallite content upon contact with the aqueous electrolyte, the films can effectively transport charges and that it is rather the high water content that harms the electronic interconnectivity within the OMIEC films. These results highlight the importance of controlling water uptake in the films to impede charge transport in n-type electrochemical devices.
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Electronic immunosensors are indispensable tools for diagnostics, particularly in scenarios demanding immediate results. Conventionally, these sensors rely on the chemical immobilization of antibodies onto electrodes. However, globular proteins tend to adsorb and unfold on these surfaces. Therefore, self-assembled monolayers (SAMs) of thiolated alkyl molecules are commonly used for indirect gold-antibody coupling. Here, a limitation associated with SAMs is revealed, wherein they curtail the longevity of protein sensors, particularly when integrated into the state-of-the-art transducer of organic bioelectronics-the organic electrochemical transistor. The SpyDirect method is introduced, generating an ultrahigh-density array of oriented nanobody receptors stably linked to the gold electrode without any SAMs. It is accomplished by directly coupling cysteine-terminated and orientation-optimized spyTag peptides, onto which nanobody-spyCatcher fusion proteins are autocatalytically attached, yielding a dense and uniform biorecognition layer. The structure-guided design optimizes the conformation and packing of flexibly tethered nanobodies. This biolayer enhances shelf-life and reduces background noise in various complex media. SpyDirect functionalization is faster and easier than SAM-based methods and does not necessitate organic solvents, rendering the sensors eco-friendly, accessible, and amenable to scalability. SpyDirect represents a broadly applicable biofunctionalization method for enhancing the cost-effectiveness, sustainability, and longevity of electronic biosensors, all without compromising sensitivity.
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Técnicas Biosensibles , Técnicas Biosensibles/métodos , Técnicas Biosensibles/instrumentación , Oro/química , Electrodos , Técnicas Electroquímicas/métodos , Técnicas Electroquímicas/instrumentación , Anticuerpos de Dominio Único/químicaRESUMEN
In Drosophila subperineurial glia (SPG) ensheath and insulate the nerve. SPG is under strict cell cycle and survival control because cell division or death of such a cell type would compromise the integrity of the blood-nerve barrier. The mechanisms underlying the survival of SPG remain unknown. Here, we show that the embryonic peripheral glia expresses the Zfh1 transcription factor, and in zfh1 mutants a particular SPG subtype, ePG10, undergoes apoptosis. Our findings show that in ePG10, Zfh1 represses the pro-apoptotic RHG-motif gene reaper in a cell-autonomous manner. Zfh1 also blocks the activation of the Jun N-terminal kinase (JNK) pathway, and reducing or enhancing JNK signalling in zfh1 mutants prevents or promotes ePG10 apoptosis. Our study shows a novel function for Zfh1 as an anti-apoptotic molecule and uncovers a cryptic JNK-dependent apoptotic programme in ePG10, which is normally blocked by Zfh1. We propose that, in cells such as SPG that do not undergo self-renewal and survive long periods, transcriptional control of RHG-motif gene expression together with fine tuning of JNK signalling is crucial for cell survival.
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Apoptosis/fisiología , Proteínas de Drosophila/fisiología , Drosophila melanogaster/fisiología , Proteínas Quinasas JNK Activadas por Mitógenos/fisiología , Neuroglía/citología , Neuroglía/metabolismo , Proteínas Represoras/fisiología , Animales , Apoptosis/genética , Western Blotting , Diferenciación Celular/genética , Diferenciación Celular/fisiología , Proteínas de Drosophila/genética , Drosophila melanogaster/enzimología , Drosophila melanogaster/metabolismo , Proteínas Inhibidoras de la Apoptosis/genética , Proteínas Inhibidoras de la Apoptosis/fisiología , Proteínas Quinasas JNK Activadas por Mitógenos/genética , Sistema Nervioso Periférico/metabolismo , Proteínas Represoras/genética , Transducción de Señal/genética , Transducción de Señal/fisiologíaRESUMEN
In alignment with widespread interest in carbon neutralization and sustainable practices, we disclose that conjugated polyelectrolyte (CPE) hydrogels are a type of recyclable, electrochemically stable, and environmentally friendly pseudocapacitive material for energy storage applications. By leveraging ionic-electronic coupling in a relatively fluid medium, one finds that hydrogels prepared using a fresh batch of an anionic CPE, namely, Pris-CPE-K, exhibit a specific capacitance of 32.6 ± 6.6 F g-1 in 2 M NaCl and are capable of 80% (26.1 ± 6.5 F g-1) capacitance retention after 100,000 galvanostatic charge-discharge (GCD) cycles at a current density (J) of 10 A g-1. We note that equilibration under a constant potential prior to GCD analysis leads to the K+ counterions in the CPE exchanging with Na+ and, thus, the relevant active material Pris-CPE-Na. It is possible to remove the CPE material from the electrochemical cell via extraction with water and to carry out a simple purification through dialysis to produce a recycled material, namely Re-CPE-Na. The recycling workup has no significant detrimental impact on the electrochemical performance. Specifically, Re-CPE-Na hydrogels display an initial specific capacitance of 26.3 ± 1.2 F g-1 (at 10 A g-1) and retain 77% of the capacitance after a subsequent 100,000 GCD cycles. Characterization by NMR, FTIR, and Raman spectroscopies, together with XPS and GPC measurements, revealed no change in the structure of the backbone or side chains. However, rheological measurements gave evidence of a slight loss in G' and G''. Overall, that CPE hydrogels display recyclability argues in favor of considering them as a novel materials platform for energy storage applications within an economically viable circular recycling strategy.
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The tight regulation of the glucose concentration in the body is crucial for balanced physiological function. We developed an electrochemical transistor comprising an n-type conjugated polymer film in contact with a catalytic enzyme for sensitive and selective glucose detection in bodily fluids. Despite the promise of these sensors, the property of the polymer that led to such high performance has remained unknown, with charge transport being the only characteristic under focus. Here, we studied the impact of the polymer chemical structure on film surface properties and enzyme adsorption behavior using a combination of physiochemical characterization methods and correlated our findings with the resulting sensor performance. We developed five n-type polymers bearing the same backbone with side chains differing in polarity and charge. We found that the nature of the side chains modulated the film surface properties, dictating the extent of interactions between the enzyme and the polymer film. Quartz crystal microbalance with dissipation monitoring studies showed that hydrophobic surfaces retained more enzymes in a densely packed arrangement, while hydrophilic surfaces captured fewer enzymes in a flattened conformation. X-ray photoelectron spectroscopy analysis of the surfaces revealed strong interactions of the enzyme with the glycolated side chains of the polymers, which improved for linear side chains compared to those for branched ones. We probed the alterations in the enzyme structure upon adsorption using circular dichroism, which suggested protein denaturation on hydrophobic surfaces. Our study concludes that a negatively charged, smooth, and hydrophilic film surface provides the best environment for enzyme adsorption with desired mass and conformation, maximizing the sensor performance. This knowledge will guide synthetic work aiming to establish close interactions between proteins and electronic materials, which is crucial for developing high-performance enzymatic metabolite biosensors and biocatalytic charge-conversion devices.
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Conjugated polymer films, which can conduct both ionic and electronic charges, are central to building soft electronic sensors and actuators. Despite the possible interplay between light absorption and the mixed conductivity of these materials in aqueous biological media, no single polymer film has been utilized to create a solar-switchable organic bioelectronic circuit that relies on a fully reversible and redox reaction-free potentiometric photodetection and current modulation. Here we demonstrate that the absorption of light by an electron and cation-transporting polymer film reversibly modulates its electrochemical potential and conductivity in an aqueous electrolyte, which is harnessed to design an n-type photo-electrochemical transistor (n-OPECT). By controlling the intensity of light incident on the n-type polymeric gate electrode, we generate transistor output characteristics that mimic the modulation of the polymeric channel current achieved through gate voltage control. The micron-scale n-OPECT exhibits a high signal-to-noise ratio and an excellent sensitivity to low light intensities. We demonstrate three direct applications of the n-OPECT, i.e., a photoplethysmogram recorder, a light-controlled inverter circuit, and a light-gated artificial synapse, underscoring the suitability of this platform for a myriad of biomedical applications that involve light intensity changes.
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Organic electrochemical transistors (OECTs) are becoming increasingly ubiquitous in various applications at the interface with biological systems. However, their widespread use is hampered by the scarcity of electron-conducting (n-type) backbones and the poor performance and stability of the existing n-OECTs. Here, we introduce organic salts as a solution additive to improve the transduction capability, shelf life, and operational stability of n-OECTs. We demonstrate that the salt-cast devices present a 10-fold increase in transconductance and achieve at least one year-long stability, while the pristine devices degrade within four months of storage. The salt-added films show improved backbone planarity and greater charge delocalization, leading to higher electronic charge carrier mobility. These films show a distinctly porous morphology where the interconnectivity is affected by the salt type, responsible for OECT speed. The salt-based films display limited changes in morphology and show lower water uptake upon electrochemical doping, a possible reason for the improved device cycling stability. Our work provides a new and easy route to improve n-type OECT performance and stability, which can be adapted for other electrochemical devices with n-type films operating at the aqueous electrolyte interface.
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Poly(3,4-ethylenedioxythiophene) (PEDOT) is a prime example of conducting polymer materials for supercapacitor electrodes that offer ease of processability and sophisticated chemical stability during operation and storage in aqueous environments. Yet, continuous improvement of its electrochemical capacitance and stability upon long cycles remains a major interest in the field, such as developing PEDOT-based composites. This work evaluates the electrochemical performances of hydroxymethyl PEDOT (PEDOTOH) coupled with hydrogel additives, namely poly(ethylene oxide) (PEO), poly(acrylic acid) (PAA), and polyethyleneimine (PEI), fabricated via a single-step electrochemical polymerization method in an aqueous solution. The PEDOTOH/PEO composite exhibits the highest capacitance (195.2â F g-1 ) compared to pristine PEDOTOH (153.9â F g-1 ), PEDOTOH/PAA (129.9â F g-1 ), and PEDOTOH/PEI (142.3â F g-1 ) at a scan rate of 10â mV s-1 . The PEDOTOH/PEO electrodes were then assembled into a symmetrical supercapacitor in an agarose gel. The type of supporting electrolytes and salt concentrations were further examined to identify the optimal agarose-based gel electrolyte. The supercapacitors comprising 2â M agarose-LiClO4 achieved a specific capacitance of 27.6â F g-1 at a current density of 2â A g-1 , a capacitance retention of â¼94% after 10,000 charge/discharge cycles at 10.6â A g-1 , delivering a maximum energy and power densities of 11.2â Wh kg-1 and 17.28â kW kg-1 , respectively. The performance of the proposed supercapacitor outperformed several reported PEDOT-based supercapacitors, including PEDOT/carbon fiber, PEDOT/CNT, and PEDOT/graphene composites. This study provides insights into the effect of incorporated hydrogel in the PEDOTOH network and the optimal conditions of agarose-based gel electrolytes for high-performance PEDOT-based supercapacitor devices.
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Electrólitos , Hidrogeles , Capacidad Eléctrica , Polimerizacion , SefarosaRESUMEN
Characterizing the molecular signature of a cell subtype leads to a better understanding of cell diversity, as this molecular data can identify new cellular markers and offer insights about cell function. Here, we describe an efficient protocol to separate a subtype of astrocytes, the Olig2-AS, from other glial cells by using a double reporter mouse approach and to determine the transcriptome profile of the Olig2-AS from the postnatal spinal cord using RNA-sequencing analysis. For complete details on the use and execution of this protocol, please refer to Ohayon et al. (2021).
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Astrocitos/citología , Citometría de Flujo/métodos , RNA-Seq/métodos , Transcriptoma/genética , Animales , Astrocitos/química , Astrocitos/metabolismo , Ratones , Ratones Transgénicos , Neuroglía/citología , Médula Espinal/citologíaRESUMEN
Astrocytes are recognized to be a heterogeneous population of cells that differ morphologically, functionally, and molecularly. Whether this heterogeneity results from generation of distinct astrocyte cell lineages, each functionally specialized to perform specific tasks, remains an open question. In this study, we used RNA sequencing analysis to determine the global transcriptome profile of the Olig2-expressing astrocyte subtype (Olig2-AS), a specific spinal astrocyte subtype that segregates early during development from Olig2 progenitors and differs from other spinal astrocytes by the expression of Olig2. We identified 245 differentially expressed genes. Among them, 135 exhibit higher levels of expression when compared with other populations of spinal astrocytes, indicating that these genes can serve as a "unique" functional signature of Olig2-AS. Among them, we identify two genes, inka2 and kcnip3, as specific molecular markers of the Olig2-AS in the P7 spinal cord. Our work thus reveals that Olig2 progenitors produce a unique spinal astrocyte subtype.