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Patterning and Cell Type Specification in the Developing CNS and PNS
Patterning and Cell Type Specification in the Developing CNS and PNS
Copyright
Contents
Contributors
I - Induction and patterning of the CNS and PNS
1 - Morphogens, patterning centers, and their mechanisms of action
1.1 General principles of morphogen gradients
1.1.1 History of the morphogen and morphogenetic field
1.1.2 How morphogen gradients pattern tissues
1.1.3 How morphogens are distributed
1.1.4 How morphogen signaling is transduced and interpreted
1.1.5 How morphogen gradients are converted into sharp boundaries
1.1.6 Summary-general principles of morphogen gradients
1.2 Local signaling centers and probable morphogens in the telencephalon
1.2.1 Early forebrain patterning
1.2.2 The RPC
1.2.3 The telencephalic roof plate and cortical hem
1.2.4 The antihem
1.3 BMPs as morphogens in telencephalic patterning
1.3.1 Performance objectives for a BMP gradient in the dorsal telencephalon
1.3.2 Midline expression and homeogenetic expansion of BMP production
1.3.3 BMP signaling gradient in the dorsal telencephalon
1.3.4 BMPs as dorsal telencephalic morphogens
1.3.5 Linear conversion of BMP signaling by cortical cells
1.3.6 Nonlinear conversion of BMP signaling by DTM cells
1.3.7 Summary-the BMP signaling gradient
1.4 FGF8 as a morphogen in telencephalic patterning
1.5 Interactions among signaling centers in telencephalic patterning
1.5.1 FGF8, Shh, and BMP signaling
1.5.2 Cross-regulation of BMP, FGF, and WNT signaling
1.5.3 Interactions of Shh, FGFs, and Gli3
1.6 Morphogens in human brain disease
1.6.1 Holoprosencephaly and Kallmann syndrome
1.6.2 Gradients in holoprosencephaly neuropathology
1.6.3 Gradients in other human brain disorders
References.
2 - Telencephalon patterning
2.1 Introduction
2.2 Telencephalon induction
2.2.1 The anterior neural ridge
2.2.2 FGF signaling
2.2.3 Wnt antagonism
2.2.4 Interactions of low Wnt with FGFs and BMPs
2.3 Overview of early telencephalic subdivisions
2.4 Establishing dorsal versus ventral domains
2.4.1 Shh and Gli3, two key players
2.4.2 Foxg1 and FGFs cooperatively promote ventral development
2.4.3 Establishing the dorsal telencephalic domain
2.4.4 Sharpening the dorsal-ventral border
2.4.5 The olfactory bulbs
2.5 Boundary structures as organizing centers and CR cell sources
2.5.1 Nomenclature of domains in the early telencephalic neuroepithelium
2.5.2 Specification of the hem and the antihem
2.5.2.1 Molecular mechanisms that act to position and specify the cortical hem
2.5.2.2 Molecular mechanisms that act to specify and position the antihem
2.5.3 Cajal-Retzius cells arise from four telencephalic boundary structures
2.5.4 Organizer functions of telencephalic boundary structures
2.5.4.1 Rostral signaling center/septum
2.5.4.1.1 Hem
2.5.4.2 Antihem
2.6 Subdividing ventral domains
2.6.1 The striatum and pallidum
2.6.2 The amygdala
2.6.3 An evolutionary perspective for how the neocortex arose
2.6.4 Lineage and fate mapping in the ventral telencephalon
2.7 Conclusions
Acknowledgments
References
3 - Area patterning of the mammalian neocortex
3.1 Introduction
3.1.1 Basic principles
3.1.2 Classic neocortical area patterning models
3.2 Indications that intrinsic mechanisms pattern the neocortical primordium
3.3 Morphogens impart position to the neocortical primordium
3.3.1 Morphogen signaling
3.3.2 Neocortical patterning by FGFs
3.3.3 Fgf8 regulates neocortical guidance of thalamic axons
3.3.4 Neocortical patterning by the cortical hem.
3.4 Patterning genes downstream of morphogen signaling
3.4.1 Emx2 and Pax6
3.4.2 Dmrt5/Dmrta2
3.4.3 Couptf1/Nr2f1
3.4.4 Sp8
3.4.5 Pbx
3.5 Do neocortical areas arise from dedicated progenitor cell pools?
3.5.1 Transcription factors known to pattern the NP appear in gradients, not domains
3.5.2 Mapping the cortical primordium with forebrain enhancers
3.6 The influence of thalamic innervation
3.6.1 Guidance of thalamocortical axons and area formation
3.6.2 Thalamic innervation determines the function of a cortical area
3.6.3 Effects of thalamocortical afferents on area size and cortical progenitor cells
3.6.4 Thalamic dependence of an area-specific feature
3.6.5 Two mechanisms united
3.7 Spontaneous activity and neocortical patterning
3.8 Conservation of patterning mechanisms among different mammalian species
3.9 Conclusions
References
4 - Patterning of thalamus
4.1 Introduction
4.2 Insights into diencephalic patterning
4.2.1 Columnar and neuromeric models
4.2.2 Morphologic segmentation of the diencephalon in the prosomeric model
4.2.3 Molecular regionalization of the diencephalon
4.2.3.1 Prosomere 1
4.2.3.2 Prosomere 2: the epithalamic domain
4.2.3.3 Prosomere 3
4.3 Prosomere 2: the thalamic domain
4.3.1 Cell lineages in the p2 alar plate
4.3.2 Signaling molecules during the initial patterning phase
4.3.2.1 Shh
4.3.2.2 Wnt
4.3.2.3 Fibroblast growth factor
4.3.3 Transcription factor control for neuronal identity
List of acronyms and abbreviations
References
5 - Midbrain patterning: polarity formation of the tectum, midbrain regionalization, and isthmus organizer
5.1 Introduction: brief description about midbrain
5.2 Tectum laminar formation
5.3 Optic tectum as a visual center for the lower vertebrate.
5.3.1 Retinotectal projection in a retinotopic manner
5.3.2 Polarity formation in the optic tectum
5.4 Development of midbrain from the mesencephalic brain vesicle
5.4.1 Transcription factors that determine the midbrain
5.4.2 Midbrain-hindbrain boundary formation
5.4.3 Diencephalon-mesencephalon boundary formation
5.4.4 Dorsoventral patterning in the midbrain
5.5 Isthmus organizer
5.5.1 Isthmus emanates organizing signal
5.5.2 Competence of the neural tube to Fgf8 signaling is determined by preexisting transcription factors
5.5.3 Intracellular signal transduction
5.5.4 How tectum and cerebellum are organized by isthmus organizing signal?
5.6 Concluding remarks
List of abbreviations of genes and molecules
List of abbreviations (general)
Glossary
References
6 - Cerebellar patterning
6.1 Introduction
6.2 Early formation of cerebellum
6.2.1 Morphogenetic aspect of first steps of cerebellar formation
6.2.2 Molecular mechanisms underlying initial formation of cerebellum
6.3 Three types of cerebellar patterning in adult mammals
6.3.1 Cerebellar anterior-posterior patterning
6.3.1.1 Lobes
6.3.1.2 Lobules (I-X)
6.3.1.3 Functional roles of lobes
6.3.2 Cerebellar medial-lateral patterning
6.3.2.1 Parasagittal zones
6.3.2.2 Parasagittal stripes
6.3.2.3 Correspondence between parasagittal zones and parasagittal stripes
6.3.2.4 Functional roles of parasagittal zones and stripes
6.3.3 Cerebellar outer-inner patterning
6.3.3.1 The molecular layer
6.3.3.2 The Purkinje cell layer
6.3.3.3 The granular layer
6.3.3.4 The white matter
6.3.3.5 The cerebellar nuclei
6.3.3.6 Roles of cerebellar outer-inner patterning
6.4 Formation of cerebellar patterning
6.4.1 Formation of cerebellar anterior-posterior patterning
6.4.1.1 Formation of lobes and lobules.
6.4.1.2 Cellular mechanisms underlying the formation of lobes and lobules
6.4.2 Formation of cerebellar medial-lateral patterning
6.4.2.1 Formation of parasagittal zones
6.4.2.2 Cellular and molecular mechanisms underlying the formation of parasagittal zones
6.4.2.3 Formation of parasagittal stripes
6.4.2.4 Critical roles of Purkinje cell birth date in the formation of embryonic and adult parasagittal stripes and parasagittal zones
6.4.3 Formation of cerebellar outer-inner patterning
6.4.3.1 Formation of the molecular layer
6.4.3.2 Formation of the Purkinje cell layer
6.4.3.3 Formation of the granular layer
6.4.3.4 Formation of the white matter and the cerebellar nuclei
6.4.3.5 Mechanisms underlying the control of neuronal migration
6.4.3.6 The deficits of neuronal migration by exposure to toxic substances and natural environmental factors result in abnormal O-I ...
References
7 - Patterning and generation of neural diversity in the spinal cord
7.1 Introduction
7.2 Spatial signals and the generation of neuronal diversity
7.2.1 Dorsoventral patterning and the induction of progenitor domains
7.2.1.1 Induction of neural progenitor ventral fate: Shh signaling
7.2.1.2 Induction of dorsal progenitor fate: Bmp and Wnt signaling
7.2.2 Rostrocaudal patterning and regional identity
7.2.2.1 Rostrocaudal antiparallel signaling
7.2.2.2 Hox function in neuronal diversity
7.3 Transcription factor combinatorial codes
7.3.1 Transcriptional codes in spinal cord progenitor fate
7.3.2 Transcription factor combinatorial codes in the diversification of postmitotic motor neurons
7.3.3 Transcriptional signatures in spinal cord interneuron diversification
7.4 Local signals and cell-cell interactions
7.4.1 The role of notchdelta signaling in interneuron and motor neuron subtype specification.
7.4.2 Retinoid signaling in motor neuron subtype specification.
Patterning and Cell Type Specification in the Developing CNS and PNS
Patterning and Cell Type Specification in the Developing CNS and PNS
Copyright
Contents
Contributors
I - Induction and patterning of the CNS and PNS
1 - Morphogens, patterning centers, and their mechanisms of action
1.1 General principles of morphogen gradients
1.1.1 History of the morphogen and morphogenetic field
1.1.2 How morphogen gradients pattern tissues
1.1.3 How morphogens are distributed
1.1.4 How morphogen signaling is transduced and interpreted
1.1.5 How morphogen gradients are converted into sharp boundaries
1.1.6 Summary-general principles of morphogen gradients
1.2 Local signaling centers and probable morphogens in the telencephalon
1.2.1 Early forebrain patterning
1.2.2 The RPC
1.2.3 The telencephalic roof plate and cortical hem
1.2.4 The antihem
1.3 BMPs as morphogens in telencephalic patterning
1.3.1 Performance objectives for a BMP gradient in the dorsal telencephalon
1.3.2 Midline expression and homeogenetic expansion of BMP production
1.3.3 BMP signaling gradient in the dorsal telencephalon
1.3.4 BMPs as dorsal telencephalic morphogens
1.3.5 Linear conversion of BMP signaling by cortical cells
1.3.6 Nonlinear conversion of BMP signaling by DTM cells
1.3.7 Summary-the BMP signaling gradient
1.4 FGF8 as a morphogen in telencephalic patterning
1.5 Interactions among signaling centers in telencephalic patterning
1.5.1 FGF8, Shh, and BMP signaling
1.5.2 Cross-regulation of BMP, FGF, and WNT signaling
1.5.3 Interactions of Shh, FGFs, and Gli3
1.6 Morphogens in human brain disease
1.6.1 Holoprosencephaly and Kallmann syndrome
1.6.2 Gradients in holoprosencephaly neuropathology
1.6.3 Gradients in other human brain disorders
References.
2 - Telencephalon patterning
2.1 Introduction
2.2 Telencephalon induction
2.2.1 The anterior neural ridge
2.2.2 FGF signaling
2.2.3 Wnt antagonism
2.2.4 Interactions of low Wnt with FGFs and BMPs
2.3 Overview of early telencephalic subdivisions
2.4 Establishing dorsal versus ventral domains
2.4.1 Shh and Gli3, two key players
2.4.2 Foxg1 and FGFs cooperatively promote ventral development
2.4.3 Establishing the dorsal telencephalic domain
2.4.4 Sharpening the dorsal-ventral border
2.4.5 The olfactory bulbs
2.5 Boundary structures as organizing centers and CR cell sources
2.5.1 Nomenclature of domains in the early telencephalic neuroepithelium
2.5.2 Specification of the hem and the antihem
2.5.2.1 Molecular mechanisms that act to position and specify the cortical hem
2.5.2.2 Molecular mechanisms that act to specify and position the antihem
2.5.3 Cajal-Retzius cells arise from four telencephalic boundary structures
2.5.4 Organizer functions of telencephalic boundary structures
2.5.4.1 Rostral signaling center/septum
2.5.4.1.1 Hem
2.5.4.2 Antihem
2.6 Subdividing ventral domains
2.6.1 The striatum and pallidum
2.6.2 The amygdala
2.6.3 An evolutionary perspective for how the neocortex arose
2.6.4 Lineage and fate mapping in the ventral telencephalon
2.7 Conclusions
Acknowledgments
References
3 - Area patterning of the mammalian neocortex
3.1 Introduction
3.1.1 Basic principles
3.1.2 Classic neocortical area patterning models
3.2 Indications that intrinsic mechanisms pattern the neocortical primordium
3.3 Morphogens impart position to the neocortical primordium
3.3.1 Morphogen signaling
3.3.2 Neocortical patterning by FGFs
3.3.3 Fgf8 regulates neocortical guidance of thalamic axons
3.3.4 Neocortical patterning by the cortical hem.
3.4 Patterning genes downstream of morphogen signaling
3.4.1 Emx2 and Pax6
3.4.2 Dmrt5/Dmrta2
3.4.3 Couptf1/Nr2f1
3.4.4 Sp8
3.4.5 Pbx
3.5 Do neocortical areas arise from dedicated progenitor cell pools?
3.5.1 Transcription factors known to pattern the NP appear in gradients, not domains
3.5.2 Mapping the cortical primordium with forebrain enhancers
3.6 The influence of thalamic innervation
3.6.1 Guidance of thalamocortical axons and area formation
3.6.2 Thalamic innervation determines the function of a cortical area
3.6.3 Effects of thalamocortical afferents on area size and cortical progenitor cells
3.6.4 Thalamic dependence of an area-specific feature
3.6.5 Two mechanisms united
3.7 Spontaneous activity and neocortical patterning
3.8 Conservation of patterning mechanisms among different mammalian species
3.9 Conclusions
References
4 - Patterning of thalamus
4.1 Introduction
4.2 Insights into diencephalic patterning
4.2.1 Columnar and neuromeric models
4.2.2 Morphologic segmentation of the diencephalon in the prosomeric model
4.2.3 Molecular regionalization of the diencephalon
4.2.3.1 Prosomere 1
4.2.3.2 Prosomere 2: the epithalamic domain
4.2.3.3 Prosomere 3
4.3 Prosomere 2: the thalamic domain
4.3.1 Cell lineages in the p2 alar plate
4.3.2 Signaling molecules during the initial patterning phase
4.3.2.1 Shh
4.3.2.2 Wnt
4.3.2.3 Fibroblast growth factor
4.3.3 Transcription factor control for neuronal identity
List of acronyms and abbreviations
References
5 - Midbrain patterning: polarity formation of the tectum, midbrain regionalization, and isthmus organizer
5.1 Introduction: brief description about midbrain
5.2 Tectum laminar formation
5.3 Optic tectum as a visual center for the lower vertebrate.
5.3.1 Retinotectal projection in a retinotopic manner
5.3.2 Polarity formation in the optic tectum
5.4 Development of midbrain from the mesencephalic brain vesicle
5.4.1 Transcription factors that determine the midbrain
5.4.2 Midbrain-hindbrain boundary formation
5.4.3 Diencephalon-mesencephalon boundary formation
5.4.4 Dorsoventral patterning in the midbrain
5.5 Isthmus organizer
5.5.1 Isthmus emanates organizing signal
5.5.2 Competence of the neural tube to Fgf8 signaling is determined by preexisting transcription factors
5.5.3 Intracellular signal transduction
5.5.4 How tectum and cerebellum are organized by isthmus organizing signal?
5.6 Concluding remarks
List of abbreviations of genes and molecules
List of abbreviations (general)
Glossary
References
6 - Cerebellar patterning
6.1 Introduction
6.2 Early formation of cerebellum
6.2.1 Morphogenetic aspect of first steps of cerebellar formation
6.2.2 Molecular mechanisms underlying initial formation of cerebellum
6.3 Three types of cerebellar patterning in adult mammals
6.3.1 Cerebellar anterior-posterior patterning
6.3.1.1 Lobes
6.3.1.2 Lobules (I-X)
6.3.1.3 Functional roles of lobes
6.3.2 Cerebellar medial-lateral patterning
6.3.2.1 Parasagittal zones
6.3.2.2 Parasagittal stripes
6.3.2.3 Correspondence between parasagittal zones and parasagittal stripes
6.3.2.4 Functional roles of parasagittal zones and stripes
6.3.3 Cerebellar outer-inner patterning
6.3.3.1 The molecular layer
6.3.3.2 The Purkinje cell layer
6.3.3.3 The granular layer
6.3.3.4 The white matter
6.3.3.5 The cerebellar nuclei
6.3.3.6 Roles of cerebellar outer-inner patterning
6.4 Formation of cerebellar patterning
6.4.1 Formation of cerebellar anterior-posterior patterning
6.4.1.1 Formation of lobes and lobules.
6.4.1.2 Cellular mechanisms underlying the formation of lobes and lobules
6.4.2 Formation of cerebellar medial-lateral patterning
6.4.2.1 Formation of parasagittal zones
6.4.2.2 Cellular and molecular mechanisms underlying the formation of parasagittal zones
6.4.2.3 Formation of parasagittal stripes
6.4.2.4 Critical roles of Purkinje cell birth date in the formation of embryonic and adult parasagittal stripes and parasagittal zones
6.4.3 Formation of cerebellar outer-inner patterning
6.4.3.1 Formation of the molecular layer
6.4.3.2 Formation of the Purkinje cell layer
6.4.3.3 Formation of the granular layer
6.4.3.4 Formation of the white matter and the cerebellar nuclei
6.4.3.5 Mechanisms underlying the control of neuronal migration
6.4.3.6 The deficits of neuronal migration by exposure to toxic substances and natural environmental factors result in abnormal O-I ...
References
7 - Patterning and generation of neural diversity in the spinal cord
7.1 Introduction
7.2 Spatial signals and the generation of neuronal diversity
7.2.1 Dorsoventral patterning and the induction of progenitor domains
7.2.1.1 Induction of neural progenitor ventral fate: Shh signaling
7.2.1.2 Induction of dorsal progenitor fate: Bmp and Wnt signaling
7.2.2 Rostrocaudal patterning and regional identity
7.2.2.1 Rostrocaudal antiparallel signaling
7.2.2.2 Hox function in neuronal diversity
7.3 Transcription factor combinatorial codes
7.3.1 Transcriptional codes in spinal cord progenitor fate
7.3.2 Transcription factor combinatorial codes in the diversification of postmitotic motor neurons
7.3.3 Transcriptional signatures in spinal cord interneuron diversification
7.4 Local signals and cell-cell interactions
7.4.1 The role of notchdelta signaling in interneuron and motor neuron subtype specification.
7.4.2 Retinoid signaling in motor neuron subtype specification.