Chapter 9. Refinement of Connectivity (Collateral Elimination/Pruning) Refinement of Synaptic Connectivity

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1 Chapter 9 Refinement of Connectivity (Collateral Elimination/Pruning) Refinement of Synaptic Connectivity Synapses are overproduced Pruning must improve function gain of correct branches loss of incorrect branches Mechanisms activity-dependent spontaneous activity (block with TTX) patterned or unpatterned intrinsic or extrinsic sensory evoked activity (block with sensory deprivation) role of competition and NMDA receptors (block with APV) Learning LTP/LTD Critical periods Homeostatic plasticity 1

2 3 Types of Refinement aka Collateral Elimination Fig 9.1 Methodology 2

3 Assay: Electrophysiology Fig 9.2 Assay: Dual Retrograde Tracer Injections Fig 9.3 3

4 Assay: Anterograde Tracer Injections Development of Ocular Dominance Columns Fig 9.4 Assay: In vivo imaging 4

5 Examples of Refinement Example: Refinement at the NMJ Polyneuronal innervation is the immature condition. In adults each muscle fiber gets input from only one motor neuron. Fig 9.6 5

6 Example: Refinement of chick retinotectal axons Fig 6.25 Example: Refinement of eye-specific laminae in LGN Retino-geniculate projection: Refinement of laminar segregation in LGN occurs in stages. Fig 9.5 6

7 Refinement of Synaptic Connectivity Synapses are overproduced Pruning must improve function gain of correct branches loss of incorrect branches Mechanisms activity-dependent spontaneous activity (block with TTX) patterned or unpatterned intrinsic or extrinsic sensory evoked activity (block with sensory deprivation) role of competition and NMDA receptors (block with APV) Learning LTP/LTD Critical periods Homeostatic plasticity The Classic Example: Development of Ocular Dominance Columns Fig 9.4 7

8 Importance of Neural Activity Activity-dependent refinement of synaptic connectivity when an axon of cell A is near enough to excite a cell B and repeatedly and consistently takes part in firing it, some growth or metabolic change takes place such that A s efficiency, as one of the cells firing B, is increased. -Donald Hebb, 1949 i.e., Fire together, Wire together! Use it or lose it! Input Layer Target 8

9 Use it or Lose it! Experience shapes brain circuitry during development. It can also be harnessed for map plasticity Map refinement: Fire together, wire together Retina Juvenile SC Retina Adult SC 9

10 Types of activity: Early patterned spontaneous activity Fig 9.21 Spontaneous calcium waves in cortex Fig

11 AChR clustering is activity-dependent Denervation Supersensitivity Fig 8.30 Synapse elimination in muscle is activity-dependent Decreased activity retards synapse elimination Increased activity accelerates synapse elimination Fig 9.13 Fig

12 Spatial separation influences synaptic competition Fig 9.15 Mechanisms of Neural Activity Effects 12

13 Heterosynaptic Depression Fig 9.26 Heterosynaptic depression at the NMJ Activity-dependent synapse elimination Red-AChR Green- unstim mn Fig

14 Synapse withdrawal mediated by pro-bdnf Growth cone collapse Muscle cell recordings Fig 9.28 Role of CAMs in mediating the effects of activity Active CAMKII promotes synapse elimination Fig

15 Role of Ca++ in synaptic depression non-coincident coincident Fig 9.29 Dendritic spine motility Effects of deprivation are age-dependent Fig

16 Activity-dependent dendritic refinement in auditory brainstem Fig 9.36 Activity-dependence of OD column development Spontaneous activity No activity Bilateral Fig

17 A Classic Example Ocular Dominance Columns in Visual Cortex Fig

18 Effects of monocular deprivation on OD columns ocular dominance plasticity Fig 9.8 Critical period for OD Plasticity Juvenile Adult Fig

19 Effects of binocular deprivation: Evidence for interocular competition Fig 9.9 Effects of strabismus/amblyopia Fig

20 Binocularity requires coincident input activity Fig 9.11 OD columns can be induced in frog tectum through competition 20

21 Mechanisms Competition mediates refinement of connectivity Fig

22 Mechanism: NMDARs 1. Mg++ block at rest 2. Requires prior depolarization 3. Passes Ca++ Developmental changes 22

23 Mechanism: NMDA Receptors act as coincidence detectors PN25091.JPG Signal Cascade 23

24 Binocular competition is mediated by NMDARs Fig 9.30 Critical Periods and the Importance of Early Experience 24

25 Critical period for OD Plasticity Juvenile Adult Fig 9.24 Early experience affects sensory processing circuitry for visual orientation and motion Fig

26 Selectivity for visual stimulus orientation is present before eye opening and is resistant to deprivation, but direction selectivity is not Fig 9.18 Early noise degrades adult auditory circuitry Fig

27 Critical period for LTD Fig 9.32 Critical Periods and GABA Fig

28 GABA is necessary for development of inhibitory connectivity Fig 9.34 Homeostatic Plasticity 28

29 Homeostatic Plasticity Fig 9.33 Development and Plasticity of Topographic Maps 29

30 Early experience affects adult auditory processing circuitry Fig 9.16 Refinement of sound localization maps requires activity Refinement of inhibitory connectivity Fig

31 Alignment of binocular maps Fig 9.19 Experience-dependence in maps of sound location Fig

32 Refinement of Synaptic Connectivity Synapses are overproduced Pruning must improve function gain of correct branches loss of incorrect branches Mechanisms activity-dependent spontaneous activity (block with TTX) patterned or unpatterned intrinsic or extrinsic sensory evoked activity (block with sensory deprivation) role of competition and NMDA receptors (block with APV) Learning LTP/LTD Critical periods Homeostatic plasticity 32

33 Example: ocular dominance columns in visual cortex are sharpened during development Fig

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