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Pyramidal cell

Pyramidal cells, or pyramidal neurons, are a type of multipolar neuron found in areas of the brain including the cerebral cortex, the hippocampus, and the amygdala. Pyramidal neurons are the primary excitation units of the mammalian prefrontal cortex and the corticospinal tract. Pyramidal neurons are also one of two cell types where the characteristic sign, Negri bodies, are found in post-mortem rabies infection.[1] Pyramidal neurons were first discovered and studied by Santiago Ramón y Cajal.[2][3] Since then, studies on pyramidal neurons have focused on topics ranging from neuroplasticity to cognition.

Pyramidal cell
A human neocortical pyramidal neuron stained via Golgi's method. The apical dendrite extends vertically above the soma (cell body) and the numerous basal dendrites radiate laterally from the base of the cell body.
A reconstruction of a pyramidal cell. Soma and dendrites are labeled in red, axon arbor in blue. (1) Soma, (2) Basal dendrite, (3) Apical dendrite, (4) Axon, (5) Collateral axon.
Details
LocationCerebral cortex esp. Layers III and V
ShapeMultipolar Pyramidal
Functionexcitatory projection neuron
NeurotransmitterGlutamate, GABA
Identifiers
MeSHD017966
NeuroLex IDsao862606388
THH1.00.01.0.00044
FMA84105
Anatomical terms of neuroanatomy
[edit on Wikidata]

Structure

One of the main structural features of the pyramidal neuron is the conic shaped soma, or cell body, after which the neuron is named. Other key structural features of the pyramidal cell are a single axon, a large apical dendrite, multiple basal dendrites, and the presence of dendritic spines.[4]

Apical dendrite

The apical dendrite rises from the apex of the pyramidal cell's soma. The apical dendrite is a single, long, thick dendrite that branches several times as distance from the soma increases and extends towards the cortical surface.[4]

Basal dendrite

Basal dendrites arise from the base of the soma. The basal dendritic tree consists of three to five primary dendrites. As distance increases from the soma, the basal dendrites branch profusely.[4]

Pyramidal cells are among the largest neurons in the brain. Both in humans and rodents, pyramidal cell bodies (somas) average around 20 μm in length. Pyramidal dendrites typically range in diameter from half a micrometer to several micrometers. The length of a single dendrite is usually several hundred micrometers. Due to branching, the total dendritic length of a pyramidal cell may reach several centimeters. The pyramidal cell's axon is often even longer and extensively branched, reaching many centimeters in total length.

Dendritic spines

Dendritic spines receive most of the excitatory impulses (EPSPs) that enter a pyramidal cell. Dendritic spines were first noted by Ramón y Cajal in 1888 by using Golgi's method. Ramón y Cajal was also the first person to propose the physiological role of increasing the receptive surface area of the neuron. The greater the pyramidal cell's surface area, the greater the neuron's ability to process and integrate large amounts of information. Dendritic spines are absent on the soma, while the number increases away from it.[3] The typical apical dendrite in a rat has at least 3,000 dendritic spines. The average human apical dendrite is approximately twice the length of a rat's, so the number of dendritic spines present on a human apical dendrite could be as high as 6,000.[5]

Growth and development

Differentiation

Pyramidal specification occurs during early development of the cerebrum. Progenitor cells are committed to the neuronal lineage in the subcortical proliferative ventricular zone (VZ) and the subventricular zone (SVZ). Immature pyramidal cells undergo migration to occupy the cortical plate, where they further diversify. Endocannabinoids (eCBs) are one class of molecules that have been shown to direct pyramidal cell development and axonal pathfinding.[6] Transcription factors such as Ctip2 and Sox5 have been shown to contribute to the direction in which pyramidal neurons direct their axons.[7]

Early postnatal development

Pyramidal cells in rats have been shown to undergo many rapid changes during early postnatal life. Between postnatal days 3 and 21, pyramidal cells have been shown to double in the size of the soma, increase in length of the apical dendrite by fivefold, and increase in basal dendrite length by thirteen-fold. Other changes include the lowering of the membrane's resting potential, reduction of membrane resistance, and an increase in the peak values of action potentials.[8]

Signaling

Like dendrites in most other neurons, the dendrites are generally the input areas of the neuron, while the axon is the neuron's output. Both axons and dendrites are highly branched. The large amount of branching allows the neuron to send and receive signals to and from many different neurons.

Pyramidal neurons, like other neurons, have numerous voltage-gated ion channels. In pyramidal cells, there is an abundance of Na+, Ca2+, and K+ channels in the dendrites, and some channels in the soma.[9][10] Ion channels within pyramidal cell dendrites have different properties from the same ion channel type within the pyramidal cell soma.[11][12] Voltage-gated Ca2+ channels in pyramidal cell dendrites are activated by subthreshold EPSPs and by back-propagating action potentials. The extent of back-propagation of action potentials within pyramidal dendrites depends upon the K+ channels. K+ channels in pyramidal cell dendrites provide a mechanism for controlling the amplitude of action potentials.[13]

The ability of pyramidal neurons to integrate information depends on the number and distribution of the synaptic inputs they receive. A single pyramidal cell receives about 30,000 excitatory inputs and 1700 inhibitory (IPSPs) inputs. Excitatory (EPSPs) inputs terminate exclusively on the dendritic spines, while inhibitory (IPSPs) inputs terminate on dendritic shafts, the soma, and even the axon. Pyramidal neurons can be excited by the neurotransmitter glutamate,[4][14] and inhibited by the neurotransmitter GABA.[4]

Firing classifications

Pyramidal neurons have been classified into different subclasses based upon their firing responses to 400-1000 millisecond current pulses. These classification are RSad, RSna, and IB neurons.

RSad

RSad pyramidal neurons, or adapting regular spiking neurons, fire with individual action potentials (APs), which are followed by a hyperpolarizing afterpotential. The afterpotential increases in duration which creates spike frequency adaptation (SFA) in the neuron.[15]

RSna

RSna pyramidal neurons, or non-adapting regular spiking neurons, fire a train of action potentials after a pulse. These neurons show no signs of adaptation.[15]

IB

IB pyramidal neurons, or intrinsically bursting neurons, respond to threshold pulses with a burst of two to five rapid action potentials. IB pyramidal neurons show no adaptation.[15]

Molecular classifications

There are several studies showing that morphological and electric pyramidal cells properties could be deduced from gene expression measured by single cell sequencing.[16] Several studies are proposing single cell classifications in mouse[17] and human[18] neurons based on gene expression could explain various neuronal properties . Neuronal types in these classifications are split into excitatory, inhibitory and hundreds of corresponding sub-bytes. For example, pyramidal cells of layer 2-3 in human are classified as FREM3 type[16] and often have high amount of Ih-current[19] generated by HCN-channel.

Function

Corticospinal tract

Pyramidal neurons are the primary neural cell type in the corticospinal tract. Normal motor control depends on the development of connections between the axons in the corticospinal tract and the spinal cord. Pyramidal cell axons follow cues such as growth factors to make specific connections. With proper connections, pyramidal cells take part in the circuitry responsible for vision guided motor function.[20]

Cognition

Pyramidal neurons in the prefrontal cortex are implicated in cognitive ability. In mammals, the complexity of pyramidal cells increases from posterior to anterior brain regions. The degree of complexity of pyramidal neurons is likely linked to the cognitive capabilities of different anthropoid species. Pyramidal cells within the prefrontal cortex appear to be responsible for processing input from the primary auditory cortex, primary somatosensory cortex, and primary visual cortex, all of which process sensory modalities[citation needed]. These cells might also play a critical role in complex object recognition within the visual processing areas of the cortex.[2]

See also

References

  1. ^ Sketchy Group, LLC. . SketchyMedical. Archived from the original on 2017-04-13.
  2. ^ a b Elston GN (November 2003). "Cortex, cognition and the cell: new insights into the pyramidal neuron and prefrontal function". Cereb. Cortex. 13 (11): 1124–38. doi:10.1093/cercor/bhg093. PMID 14576205.
  3. ^ a b García-López P, García-Marín V, Freire M (November 2006). "Three-dimensional reconstruction and quantitative study of a pyramidal cell of a Cajal histological preparation". J. Neurosci. 26 (44): 11249–52. doi:10.1523/JNEUROSCI.3543-06.2006. PMC 6674523. PMID 17079652.
  4. ^ a b c d e Megías M, Emri Z, Freund TF, Gulyás AI (2001). "Total number and distribution of inhibitory and excitatory synapses on hippocampal CA1 pyramidal cells". Neuroscience. 102 (3): 527–40. doi:10.1016/S0306-4522(00)00496-6. PMID 11226691. S2CID 16458290.
  5. ^ Laberge D, Kasevich R (November 2007). "The apical dendrite theory of consciousness". Neural Netw. 20 (9): 1004–20. doi:10.1016/j.neunet.2007.09.006. PMID 17920812.
  6. ^ Mulder J, Aguado T, Keimpema E, et al. (June 2008). "Endocannabinoid signaling controls pyramidal cell specification and long-range axon patterning". Proc. Natl. Acad. Sci. U.S.A. 105 (25): 8760–5. Bibcode:2008PNAS..105.8760M. doi:10.1073/pnas.0803545105. PMC 2438381. PMID 18562289.
  7. ^ Fishell G, Hanashima C (February 2008). "Pyramidal neurons grow up and change their mind". Neuron. 57 (3): 333–8. doi:10.1016/j.neuron.2008.01.018. PMID 18255026. S2CID 15095100.
  8. ^ Zhang ZW (March 2004). "Maturation of layer V pyramidal neurons in the rat prefrontal cortex: intrinsic properties and synaptic function". J. Neurophysiol. 91 (3): 1171–82. doi:10.1152/jn.00855.2003. PMID 14602839.
  9. ^ Spruston, Nelson (2008). "Pyramidal neurons: dendritic structure and synaptic integration". Nature Reviews Neuroscience. 9 (3): 206–221. doi:10.1038/nrn2286. PMID 18270515. S2CID 1142249.
  10. ^ Georgiev, Danko D.; Kolev, Stefan K.; Cohen, Eliahu; Glazebrook, James F. (2020). "Computational capacity of pyramidal neurons in the cerebral cortex". Brain Research. 1748: 147069. arXiv:2009.10615. doi:10.1016/j.brainres.2020.147069. PMID 32858030. S2CID 221277603.
  11. ^ Golding, Nace L.; Mickus, Timothy J.; Katz, Yael; Kath, William L.; Spruston, Nelson (2005). "Factors mediating powerful voltage attenuation along CA1 pyramidal neuron dendrites". Journal of Physiology. 568 (1): 69–82. doi:10.1113/jphysiol.2005.086793. PMC 1474764. PMID 16002454.
  12. ^ Remy, S.; Beck, H.; Yaari, Y. (2010). "Plasticity of voltage-gated ion channels in pyramidal cell dendrites". Current Opinion in Neurobiology. 20 (4): 503–509. doi:10.1016/j.conb.2010.06.006. PMID 20691582. S2CID 4713853.
  13. ^ Magee J, Hoffman D, Colbert C, Johnston D (1998). "Electrical and calcium signaling in dendrites of hippocampal pyramidal neurons". Annu. Rev. Physiol. 60 (1): 327–46. doi:10.1146/annurev.physiol.60.1.327. PMID 9558467.
  14. ^ Wong, R. K. S.; Traub, R. D. (2009-01-01), "NETWORKS | Cellular Properties and Synaptic Connectivity of CA3 Pyramidal Cells: Mechanisms for Epileptic Synchronization and Epileptogenesis", in Schwartzkroin, Philip A. (ed.), Encyclopedia of Basic Epilepsy Research, Oxford: Academic Press, pp. 815–819, doi:10.1016/b978-012373961-2.00215-0, ISBN 978-0-12-373961-2, retrieved 2020-11-18
  15. ^ a b c Franceschetti S, Sancini G, Panzica F, Radici C, Avanzini G (April 1998). "Postnatal differentiation of firing properties and morphological characteristics in layer V pyramidal neurons of the sensorimotor cortex". Neuroscience. 83 (4): 1013–24. doi:10.1016/S0306-4522(97)00463-6. PMID 9502243. S2CID 6986307.
  16. ^ a b Berg, Jim; Sorensen, Staci A.; Ting, Jonathan T.; Miller, Jeremy A.; Chartrand, Thomas; Buchin, Anatoly; Bakken, Trygve E.; Budzillo, Agata; Dee, Nick; Ding, Song-Lin; Gouwens, Nathan W. (October 2021). "Human neocortical expansion involves glutamatergic neuron diversification". Nature. 598 (7879): 151–158. doi:10.1038/s41586-021-03813-8. ISSN 1476-4687. PMC 8494638. PMID 34616067.
  17. ^ Gouwens, Nathan W.; Sorensen, Staci A.; Berg, Jim; Lee, Changkyu; Jarsky, Tim; Ting, Jonathan; Sunkin, Susan M.; Feng, David; Anastassiou, Costas A.; Barkan, Eliza; Bickley, Kris (July 2019). "Classification of electrophysiological and morphological neuron types in the mouse visual cortex". Nature Neuroscience. 22 (7): 1182–1195. doi:10.1038/s41593-019-0417-0. ISSN 1546-1726. PMC 8078853. PMID 31209381.
  18. ^ Bakken, Trygve E.; Jorstad, Nikolas L.; Hu, Qiwen; Lake, Blue B.; Tian, Wei; Kalmbach, Brian E.; Crow, Megan; Hodge, Rebecca D.; Krienen, Fenna M.; Sorensen, Staci A.; Eggermont, Jeroen (October 2021). "Comparative cellular analysis of motor cortex in human, marmoset and mouse". Nature. 598 (7879): 111–119. doi:10.1038/s41586-021-03465-8. ISSN 1476-4687. PMC 8494640. PMID 34616062.
  19. ^ Kalmbach, Brian E.; Buchin, Anatoly; Long, Brian; Close, Jennie; Nandi, Anirban; Miller, Jeremy A.; Bakken, Trygve E.; Hodge, Rebecca D.; Chong, Peter; de Frates, Rebecca; Dai, Kael (2018-12-05). "h-Channels Contribute to Divergent Intrinsic Membrane Properties of Supragranular Pyramidal Neurons in Human versus Mouse Cerebral Cortex". Neuron. 100 (5): 1194–1208.e5. doi:10.1016/j.neuron.2018.10.012. ISSN 0896-6273. PMID 30392798. S2CID 53218514.
  20. ^ Salimi I, Friel KM, Martin JH (July 2008). "Pyramidal tract stimulation restores normal corticospinal tract connections and visuomotor skill after early postnatal motor cortex activity blockade". J. Neurosci. 28 (29): 7426–34. doi:10.1523/JNEUROSCI.1078-08.2008. PMC 2567132. PMID 18632946.

External links

  • Pyramidal cell - Cell Centered Database
  • Image
  • Diagram (as part of slideshow) 2016-11-02 at the Wayback Machine

pyramidal, cell, pyramidal, neurons, type, multipolar, neuron, found, areas, brain, including, cerebral, cortex, hippocampus, amygdala, pyramidal, neurons, primary, excitation, units, mammalian, prefrontal, cortex, corticospinal, tract, pyramidal, neurons, als. Pyramidal cells or pyramidal neurons are a type of multipolar neuron found in areas of the brain including the cerebral cortex the hippocampus and the amygdala Pyramidal neurons are the primary excitation units of the mammalian prefrontal cortex and the corticospinal tract Pyramidal neurons are also one of two cell types where the characteristic sign Negri bodies are found in post mortem rabies infection 1 Pyramidal neurons were first discovered and studied by Santiago Ramon y Cajal 2 3 Since then studies on pyramidal neurons have focused on topics ranging from neuroplasticity to cognition Pyramidal cellA human neocortical pyramidal neuron stained via Golgi s method The apical dendrite extends vertically above the soma cell body and the numerous basal dendrites radiate laterally from the base of the cell body A reconstruction of a pyramidal cell Soma and dendrites are labeled in red axon arbor in blue 1 Soma 2 Basal dendrite 3 Apical dendrite 4 Axon 5 Collateral axon DetailsLocationCerebral cortex esp Layers III and VShapeMultipolar PyramidalFunctionexcitatory projection neuronNeurotransmitterGlutamate GABAIdentifiersMeSHD017966NeuroLex IDsao862606388THH1 00 01 0 00044FMA84105Anatomical terms of neuroanatomy edit on Wikidata Contents 1 Structure 1 1 Apical dendrite 1 2 Basal dendrite 1 3 Dendritic spines 2 Growth and development 2 1 Differentiation 2 2 Early postnatal development 3 Signaling 3 1 Firing classifications 3 1 1 RSad 3 1 2 RSna 3 1 3 IB 3 2 Molecular classifications 4 Function 4 1 Corticospinal tract 4 2 Cognition 5 See also 6 References 7 External linksStructure Edit Pyramidal neuron visualized by green fluorescent protein gfp A hippocampal pyramidal cellOne of the main structural features of the pyramidal neuron is the conic shaped soma or cell body after which the neuron is named Other key structural features of the pyramidal cell are a single axon a large apical dendrite multiple basal dendrites and the presence of dendritic spines 4 Apical dendrite Edit The apical dendrite rises from the apex of the pyramidal cell s soma The apical dendrite is a single long thick dendrite that branches several times as distance from the soma increases and extends towards the cortical surface 4 Basal dendrite Edit Basal dendrites arise from the base of the soma The basal dendritic tree consists of three to five primary dendrites As distance increases from the soma the basal dendrites branch profusely 4 Pyramidal cells are among the largest neurons in the brain Both in humans and rodents pyramidal cell bodies somas average around 20 mm in length Pyramidal dendrites typically range in diameter from half a micrometer to several micrometers The length of a single dendrite is usually several hundred micrometers Due to branching the total dendritic length of a pyramidal cell may reach several centimeters The pyramidal cell s axon is often even longer and extensively branched reaching many centimeters in total length Dendritic spines Edit Dendritic spines receive most of the excitatory impulses EPSPs that enter a pyramidal cell Dendritic spines were first noted by Ramon y Cajal in 1888 by using Golgi s method Ramon y Cajal was also the first person to propose the physiological role of increasing the receptive surface area of the neuron The greater the pyramidal cell s surface area the greater the neuron s ability to process and integrate large amounts of information Dendritic spines are absent on the soma while the number increases away from it 3 The typical apical dendrite in a rat has at least 3 000 dendritic spines The average human apical dendrite is approximately twice the length of a rat s so the number of dendritic spines present on a human apical dendrite could be as high as 6 000 5 Growth and development EditDifferentiation Edit Pyramidal specification occurs during early development of the cerebrum Progenitor cells are committed to the neuronal lineage in the subcortical proliferative ventricular zone VZ and the subventricular zone SVZ Immature pyramidal cells undergo migration to occupy the cortical plate where they further diversify Endocannabinoids eCBs are one class of molecules that have been shown to direct pyramidal cell development and axonal pathfinding 6 Transcription factors such as Ctip2 and Sox5 have been shown to contribute to the direction in which pyramidal neurons direct their axons 7 Early postnatal development Edit Pyramidal cells in rats have been shown to undergo many rapid changes during early postnatal life Between postnatal days 3 and 21 pyramidal cells have been shown to double in the size of the soma increase in length of the apical dendrite by fivefold and increase in basal dendrite length by thirteen fold Other changes include the lowering of the membrane s resting potential reduction of membrane resistance and an increase in the peak values of action potentials 8 Signaling EditLike dendrites in most other neurons the dendrites are generally the input areas of the neuron while the axon is the neuron s output Both axons and dendrites are highly branched The large amount of branching allows the neuron to send and receive signals to and from many different neurons Pyramidal neurons like other neurons have numerous voltage gated ion channels In pyramidal cells there is an abundance of Na Ca2 and K channels in the dendrites and some channels in the soma 9 10 Ion channels within pyramidal cell dendrites have different properties from the same ion channel type within the pyramidal cell soma 11 12 Voltage gated Ca2 channels in pyramidal cell dendrites are activated by subthreshold EPSPs and by back propagating action potentials The extent of back propagation of action potentials within pyramidal dendrites depends upon the K channels K channels in pyramidal cell dendrites provide a mechanism for controlling the amplitude of action potentials 13 The ability of pyramidal neurons to integrate information depends on the number and distribution of the synaptic inputs they receive A single pyramidal cell receives about 30 000 excitatory inputs and 1700 inhibitory IPSPs inputs Excitatory EPSPs inputs terminate exclusively on the dendritic spines while inhibitory IPSPs inputs terminate on dendritic shafts the soma and even the axon Pyramidal neurons can be excited by the neurotransmitter glutamate 4 14 and inhibited by the neurotransmitter GABA 4 Firing classifications Edit Pyramidal neurons have been classified into different subclasses based upon their firing responses to 400 1000 millisecond current pulses These classification are RSad RSna and IB neurons RSad Edit RSad pyramidal neurons or adapting regular spiking neurons fire with individual action potentials APs which are followed by a hyperpolarizing afterpotential The afterpotential increases in duration which creates spike frequency adaptation SFA in the neuron 15 RSna Edit RSna pyramidal neurons or non adapting regular spiking neurons fire a train of action potentials after a pulse These neurons show no signs of adaptation 15 IB Edit IB pyramidal neurons or intrinsically bursting neurons respond to threshold pulses with a burst of two to five rapid action potentials IB pyramidal neurons show no adaptation 15 Molecular classifications Edit There are several studies showing that morphological and electric pyramidal cells properties could be deduced from gene expression measured by single cell sequencing 16 Several studies are proposing single cell classifications in mouse 17 and human 18 neurons based on gene expression could explain various neuronal properties Neuronal types in these classifications are split into excitatory inhibitory and hundreds of corresponding sub bytes For example pyramidal cells of layer 2 3 in human are classified as FREM3 type 16 and often have high amount of Ih current 19 generated by HCN channel Function EditCorticospinal tract Edit Pyramidal neurons are the primary neural cell type in the corticospinal tract Normal motor control depends on the development of connections between the axons in the corticospinal tract and the spinal cord Pyramidal cell axons follow cues such as growth factors to make specific connections With proper connections pyramidal cells take part in the circuitry responsible for vision guided motor function 20 Cognition Edit Pyramidal neurons in the prefrontal cortex are implicated in cognitive ability In mammals the complexity of pyramidal cells increases from posterior to anterior brain regions The degree of complexity of pyramidal neurons is likely linked to the cognitive capabilities of different anthropoid species Pyramidal cells within the prefrontal cortex appear to be responsible for processing input from the primary auditory cortex primary somatosensory cortex and primary visual cortex all of which process sensory modalities citation needed These cells might also play a critical role in complex object recognition within the visual processing areas of the cortex 2 See also EditCerebral cortex Pyramidal tract Chandelier cells innervate initial segments of pyramidal axons Rosehip neuronReferences Edit Sketchy Group LLC 2 3 rhabdovirus SketchyMedical Archived from the original on 2017 04 13 a b Elston GN November 2003 Cortex cognition and the cell new insights into the pyramidal neuron and prefrontal function Cereb Cortex 13 11 1124 38 doi 10 1093 cercor bhg093 PMID 14576205 a b Garcia Lopez P Garcia Marin V Freire M November 2006 Three dimensional reconstruction and quantitative study of a pyramidal cell of a Cajal histological preparation J Neurosci 26 44 11249 52 doi 10 1523 JNEUROSCI 3543 06 2006 PMC 6674523 PMID 17079652 a b c d e Megias M Emri Z Freund TF Gulyas AI 2001 Total number and distribution of inhibitory and excitatory synapses on hippocampal CA1 pyramidal cells Neuroscience 102 3 527 40 doi 10 1016 S0306 4522 00 00496 6 PMID 11226691 S2CID 16458290 Laberge D Kasevich R November 2007 The apical dendrite theory of consciousness Neural Netw 20 9 1004 20 doi 10 1016 j neunet 2007 09 006 PMID 17920812 Mulder J Aguado T Keimpema E et al June 2008 Endocannabinoid signaling controls pyramidal cell specification and long range axon patterning Proc Natl Acad Sci U S A 105 25 8760 5 Bibcode 2008PNAS 105 8760M doi 10 1073 pnas 0803545105 PMC 2438381 PMID 18562289 Fishell G Hanashima C February 2008 Pyramidal neurons grow up and change their mind Neuron 57 3 333 8 doi 10 1016 j neuron 2008 01 018 PMID 18255026 S2CID 15095100 Zhang ZW March 2004 Maturation of layer V pyramidal neurons in the rat prefrontal cortex intrinsic properties and synaptic function J Neurophysiol 91 3 1171 82 doi 10 1152 jn 00855 2003 PMID 14602839 Spruston Nelson 2008 Pyramidal neurons dendritic structure and synaptic integration Nature Reviews Neuroscience 9 3 206 221 doi 10 1038 nrn2286 PMID 18270515 S2CID 1142249 Georgiev Danko D Kolev Stefan K Cohen Eliahu Glazebrook James F 2020 Computational capacity of pyramidal neurons in the cerebral cortex Brain Research 1748 147069 arXiv 2009 10615 doi 10 1016 j brainres 2020 147069 PMID 32858030 S2CID 221277603 Golding Nace L Mickus Timothy J Katz Yael Kath William L Spruston Nelson 2005 Factors mediating powerful voltage attenuation along CA1 pyramidal neuron dendrites Journal of Physiology 568 1 69 82 doi 10 1113 jphysiol 2005 086793 PMC 1474764 PMID 16002454 Remy S Beck H Yaari Y 2010 Plasticity of voltage gated ion channels in pyramidal cell dendrites Current Opinion in Neurobiology 20 4 503 509 doi 10 1016 j conb 2010 06 006 PMID 20691582 S2CID 4713853 Magee J Hoffman D Colbert C Johnston D 1998 Electrical and calcium signaling in dendrites of hippocampal pyramidal neurons Annu Rev Physiol 60 1 327 46 doi 10 1146 annurev physiol 60 1 327 PMID 9558467 Wong R K S Traub R D 2009 01 01 NETWORKS Cellular Properties and Synaptic Connectivity of CA3 Pyramidal Cells Mechanisms for Epileptic Synchronization and Epileptogenesis in Schwartzkroin Philip A ed Encyclopedia of Basic Epilepsy Research Oxford Academic Press pp 815 819 doi 10 1016 b978 012373961 2 00215 0 ISBN 978 0 12 373961 2 retrieved 2020 11 18 a b c Franceschetti S Sancini G Panzica F Radici C Avanzini G April 1998 Postnatal differentiation of firing properties and morphological characteristics in layer V pyramidal neurons of the sensorimotor cortex Neuroscience 83 4 1013 24 doi 10 1016 S0306 4522 97 00463 6 PMID 9502243 S2CID 6986307 a b Berg Jim Sorensen Staci A Ting Jonathan T Miller Jeremy A Chartrand Thomas Buchin Anatoly Bakken Trygve E Budzillo Agata Dee Nick Ding Song Lin Gouwens Nathan W October 2021 Human neocortical expansion involves glutamatergic neuron diversification Nature 598 7879 151 158 doi 10 1038 s41586 021 03813 8 ISSN 1476 4687 PMC 8494638 PMID 34616067 Gouwens Nathan W Sorensen Staci A Berg Jim Lee Changkyu Jarsky Tim Ting Jonathan Sunkin Susan M Feng David Anastassiou Costas A Barkan Eliza Bickley Kris July 2019 Classification of electrophysiological and morphological neuron types in the mouse visual cortex Nature Neuroscience 22 7 1182 1195 doi 10 1038 s41593 019 0417 0 ISSN 1546 1726 PMC 8078853 PMID 31209381 Bakken Trygve E Jorstad Nikolas L Hu Qiwen Lake Blue B Tian Wei Kalmbach Brian E Crow Megan Hodge Rebecca D Krienen Fenna M Sorensen Staci A Eggermont Jeroen October 2021 Comparative cellular analysis of motor cortex in human marmoset and mouse Nature 598 7879 111 119 doi 10 1038 s41586 021 03465 8 ISSN 1476 4687 PMC 8494640 PMID 34616062 Kalmbach Brian E Buchin Anatoly Long Brian Close Jennie Nandi Anirban Miller Jeremy A Bakken Trygve E Hodge Rebecca D Chong Peter de Frates Rebecca Dai Kael 2018 12 05 h Channels Contribute to Divergent Intrinsic Membrane Properties of Supragranular Pyramidal Neurons in Human versus Mouse Cerebral Cortex Neuron 100 5 1194 1208 e5 doi 10 1016 j neuron 2018 10 012 ISSN 0896 6273 PMID 30392798 S2CID 53218514 Salimi I Friel KM Martin JH July 2008 Pyramidal tract stimulation restores normal corticospinal tract connections and visuomotor skill after early postnatal motor cortex activity blockade J Neurosci 28 29 7426 34 doi 10 1523 JNEUROSCI 1078 08 2008 PMC 2567132 PMID 18632946 External links EditPyramidal cell Cell Centered Database Diagram Image Diagram as part of slideshow Archived 2016 11 02 at the Wayback Machine Retrieved from https en wikipedia org w index php title Pyramidal cell amp oldid 1136193812, wikipedia, wiki, book, books, library,

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