Cerebellum: Origins and Development
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1 Cerebellum: Origins and Development Found in all vertebrates Dorsal lip of developing medulla (rhombencephalon) Near terminations of vestibular (VIII) and lateral line afferents, which sense fluid displacement Also inputs from tectum and spinal cord Ancestral role: representing an image of the relations of the body in space (Bullock 1969) Self vs. non self? Deficits associated with the cerebellum Acute ablation (Rolando, Flourens, Holmes) exaggerated reflexes, ballistic movements Congenital reduction ataxia, but also autism (Courchesne) Sensory function may also be an ancestral principle 1
2 Functional roles for the cerebellum Sensory surprise / error signals Motor coordination Sensory teaching signal Enlargement of the cerebellum for echolocation and electrolocation Clark et al. (2001) Nieuwenhuys et al. 2
3 Courchesne 1997 Palmen SJMC et al. (2004) Brain, 127:
4 Timing based theories Eccles dynamic loop control hypothesis Lead lag compensator Transforming sensory into muscle space (Pellionisz and Llinas) Modulate, accelerate or synchronize movements (Llinas/Welsh/Bloedel/Thach) Smith predictor /inverse dynamics (Miall) Learning based theories The cerebellum as a site of learning (Marr/Albus/Ito) Vestibulo ocular reflex gain adaptation (Lisberger/Raymond) Eyeblink conditioning (Thompson/Mauk) Sensory expectation subtraction Adaptive learning /inverse dynamics (Miall) Subtraction of sensory expectations (Bell) Raymond, Mauk and Lisberger (1996) Science 4
5 Cerebellar Structure and Anatomy Common features of all cerebella 1. Cerebellar cortex Purkinje neurons, the sole output: inhibitory 2. Inputs: Mossy fibers>>granule cells>parallel fibers The inferior olive>climbing fibers 3. Output: The vestibular nuclei and deep nuclei 4. A few local inhibitory neurons: stellate and Golgi neurons 5
6 JC Eccles (1969) Naturwissenschaften 56: Innovations during cerebellar evolution 1. Increasing divergence of the mossy fiber pathway 2. Proliferation of inhibitory neurons: Golgi neurons (not quite the same in dogfish), basket cells (birds and mammals) 3. Lugaro cells, unipolar brush cells 6
7 Median cell counts in vertebrate cerebellar structures From data summarized in KWT Caddy and TJ Biscoe (1978) Phil Trans Roy Soc Lond Ser B 287: , with additional measurements from BB Gould and P Rakic (1981) Exp Brain Res 44: , R Alvarez and R Anadón (1987) J Hirnforsch 28: , RJ Harvey and RMA Napper(1991) Prog Neurobiol 36: , JA Heckroth (1994) J Comp Neurol 343: , and R Alvarez-Otero et al (1996) J Comp Neurol 368: Climbing fibers Powerful input causing massive dendritic spike More focused in their branching than mossy fibers Instructive signal conveying errors or other sensory events (Marr, Albus, Ito) Regulation of Purkinje cell excitability: Synchronize sodium spike activity of Purkinje cells (Llinás, Welsh) Fire together to accelerate movement (Bloedel) Kreitzer et al
8 Functions of climbing fiber firing and complex spikes Sensory events such as error signals (Simpson et al 1996 Behav Brain Sci) Instructive signal for learning (Raymond et al 1996 Science) Synchronization guides timing and activity (Welsh 2002 NYAS) Floccular target neurons in the medial vestibular nucleus: deep-nuclear homologs C Sekirnjak et al. (2003) J. Neurosci. 23:
9 Deep nuclei Output relay of cerebellum; target of Purkinje cell axon output Inhibit inferior olive and send mossy fibers back into cerebellum Possible site of learning Reverberating activity? a black box. Apps and Garwicz (2005) Nat Rev Neurosci 6: Kandel et al / Barlow 9
10 Sugihara I, Wu H-S, Shinoda Y (2001) J. Neurosci. 21: Jinno, Jeromin, and Kosaka (2004) Cerebellum 3:
11 AAV infected cells expressing D3cpv under Tet-promoter controlled by tta under human synapsin promoter Field of view 510 µm Neighboring PC dendrites show strong correlations 11
12 Calcium responses in Purkinje cell dendrites Sensory responses show intraburst synchrony Eyeblink conditioning: order-dependent associative learning 12
13 Timing-dependence of eyeblink conditioning in vivo In eyeblink conditioning, tone must precede airpuff by >80 msec I. Goremezano, E.J. Kehoe, and B.S. Marshall (1983) Prog. Psychobiol. and Physiol. Psych., 10: Cerebellar long-term depression is order-dependent Wang, Denk and Hausser (2000) Nature Neuroscience 3:
14 The IP 3 receptor: a synaptic coincidence detector Parallel fiber (IP 3 ) and climbing fiber (Ca ++ entry) activity co-activate calcium release Functions of the olivary system Cerebellar cortex: local chemical signaling Inferior olive: local and regional synchrony Deep nuclei: coincidence-based readout mechanism 14
15 Bower JM and Woolston DC (1983) J. Neurophysiol. 49: Interneuron activation in a running mouse 15
16 Interneuron activation in a running mouse 16
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Organization of The Nervous System PROF. SAEED ABUEL MAKAREM Objectives By the end of the lecture, you should be able to: List the parts of the nervous system. List the function of the nervous system.
Basal Nuclei (Ganglia)
Doctor said he will not go deep within these slides because we will take them in physiology, so he will explain the anatomical structures, and he will go faster in the functions sheet in yellow Basal Nuclei