Circuits of the central nervous system

INPUT
ELEMENTS OF NEURAL CIRCUITS ventral horn of the spinal cord Cerebral neocortex
Neural circuits process sensory information, generate ●afferent sensory axons in the ●relay neurons of the thalamus
motor output, and create spontaneous activity dorsal roots- major input to the send axons into layers of the cortex
Circuits- synaptically interconnected networks, each neuron within a circuit spinal cord; carry information from to bring a range of info about
may have very specialized properties (in complex mammals) somatic sensory receptors in the skin, sensation, motor systems, and the
●Some neural circuits may be primarily sensory (e.g., the retina) or connective tissue, and muscles body’s internal state.
●motor (e.g., the ventral horns of the spinal cord). ●local circuits in the spinal cord have ●most numerous type of input to the
●Many circuits combine features of both, with some neurons dedicated to other sources of input, including local circuits of the neocortex comes
●providing and processing sensory input, others to commanding descending input from the brain and from the neocortex —from adjacent
●motor output, and many neurons (perhaps most) doing both. input from the spinal cord, local circuits, distant areas
●Neural circuits may also generate their own intrinsic signals, with no need both from the contralateral side and of cortex, and the contralateral
for any sensory or central input to activate them. from spinal segments hemisphere.
Pacemakers – generates coordination of rhythmic temporal patterns across local circuits receive multiple types of input.
hundreds of outputs, patterns and rhythms can always be modulated—
stopped, started, or altered—by input from sensory or central pathways. OUTPUT -achieved w/ a subset of cells known as projection neurons, or
central pattern generators - Neuronal circuits that produce rhythmic motor principal neurons - send axons to one or more targets.
output ventral horn of the spinal cord Cerebral neocortex
α motor neurons – Spinal output; large pyramidal neurons in layer V-
Nervous systems have several levels of organization send their axons out through the Output axons from the neocortex;
neural subsystems and pathways - @highest level; include the sensory ventral roots to innervate skeletal innervate targets in the brainstem,
input from the retina leading to the visual cortex, the central processing muscle fibers. spinal cord & neurons in layer VI,
regions that make sense of the visual information and the motor systems that which make their synapses
coordinate movement of the eyes and head. back onto the cells of the thalamus.
information sent in both directions along sensory/central motor pathways most local circuits have multiple types of outputs.
local circuit - @cellular level, local brain region, the arrangement of neurons **neural circuit that has only input and output cells.
and their synaptic connections; includes the set of inputs, outputs, and all the Local processing - achieved by interneurons or intrinsic neurons, whose
interconnected neurons that are essential to functions of the local axonal connections remain within the local circuit.
brain region, modular in interchangeability, interconnected. interneurons or intrinsic neurons – vary in structure, function, types.
Microcircuits –W/in local circuits; finer arrangements of neurons & *Both the spinal cord & neocortex have excitatory & inhibitory interneurons
synapses interneurons or intrinsic neurons
repeated numerous times w/in a circuit, determine the transformations of info make specific or widely divergent connections, either receive direct contact
w/in dendrites & synapses. from input axons or process only information from other interneurons
**@brain: MORE interneurons than output neurons
Most local circuits have three elements:
input axons, interneurons, and projection (output) neurons “principles” of local circuits (interactive, interdependent & wide network)
the ventral horn of the spinal cord and the cerebral neocortex. ●projection cell may have some of the characteristics of an interneuron
input, a set of axons that originate elsewhere and terminate ●some interneurons may entirely lack an axon and instead make their local
in synapses within the local circuit. synaptic connections through short neurites or even dendrites.
●Rare case: source of input to a local circuit may not be purely synaptic but
chemical
 *main neurons in local circuits – wired in series, massive in numbers, operate inhibit the α motor neurons of the antagonists: reciprocal innervation
in parallel with one another, demonstrate cross talk(info is shared mutually)
SIMPLE, STEREOTYPED RESPONSES: SPINAL REFLEX CIRCUITS
Passive stretching of a skeletal muscle causes a reflexive contraction of
that same muscle and relaxation of the antagonist muscles
Reflexes- basic neural functions & involve simple neuronal circuits, essential
elements of behavior
Motor reflex - rapid, stereotyped motor response to a particular sensory
stimulus. The CNS commands the body to move about by activating motor
neurons, which excite skeletal muscles
muscle stretch- simple sensory stimulus, only one synapse
motor unit- collective term for a single motor neuron and the muscle cells
that it synapses on, each muscle cell- belong to only 1 motor unit (1:1)
size of motor units - depends on muscle function| 2 types of motor neurons:
α motor neurons - innervate the γ motor neurons innervate only the
main force-generating muscle fibers fibers of the muscle spindles Force applied to Golgi tendon organ regulate muscle contractile strength
(the extrafusal fibers) Golgi tendon organ - mechanosensory transducer; alight in series w/ the
motor neuron pool- group of all motor neurons innervating a single muscle, sensitive to tension w/in a tendon, responds to the force generated by
muscle the muscle rather than muscle length; may respond during passive muscle
stretch reflex or myotatic reflex - when a skeletal muscle is abruptly stretch, also during active contractions of a muscle
stretched, a rapid, reflexive contraction of the same muscle often occurs. The group Ib sensory axons of the tendon organs excite both excitatory and
contraction increases muscle tension and opposes the stretch; strong in inhibitory interneurons within the spinal cord.
extensor muscles(resist gravity); aka Myotatic reflex – specific for the same In some cases, this interneuron circuitry inhibits the muscle in which tension
muscle that is stretched. [also seen in biceps of the arm and the muscles that has increased and excites the antagonistic muscle; therefore, activity in the
close the jaw] | Knee jerk reflex: another ex, elicited by a light tap on the tendon organs can yield effects that are almost the opposite of the stretch
patellar tendon. The tap deflects the tendon, which then pulls on and briefly reflex. Under other circumstances, particularly during rapid movements such
stretches the quadriceps femoris muscle. A reflexive contraction of the as locomotion, sensory input from Golgi tendon organs excites the motor
quadriceps quickly follows. neurons activating the same muscle. The reflex effects of Golgi tendon organ
●depends on the nervous system & requires sensory feedback from the activity vary because the interneurons receiving input from Ib axons also
muscle| ● muscle spindles -basic circuit for the stretch reflex begins with the receive input from other sensory endings in the muscle and skin, and from
primary sensory axons from muscle spindles axons descending from the brain. In general, reflexes mediated by the Golgi
●Increasing the length of the muscle stimulates large Ia axons from the tendon organs serve to control the force within muscles and the
primary sensory endings, group Ia sensory axons terminate monosynaptically stability of joints.
onto the α motor neurons that innervate the same muscle from which the
group Ia axons originated. Noxious stimuli can evoke complex reflexive movements
**muscle stretching causes rapid feedback excitation of the same muscle flexion-withdrawal reflex: bilateral flexor reflex, both inhibitory &
through 1 sensory neuron, 1 central synapse, 1 motor neuron. excitatory. The original stimulus for the reflex came from fast pain afferent
● Monosynaptic connections -rapid component of the stretch reflex neurons in the skin- group Aδ axons.; needs circuitry on both ipsilateral and
*stretched muscle is contracting  parallel circuits are inhibiting the α motor contralateral side,
neurons of its antagonist muscles. (knee-jerk reflex causes contraction of the >specificity - withdrawal of the part in the direction opposite the side of the
quadriceps muscle, it simultaneously causes relaxation of its antagonists, stimulus
including the semitendinosus muscle) > strength of the reflex is related to the intensity of the stimulus.
** for inhibition: group Ia sensory axons excite specific interneurons that * flexor reflexes coordinate the movement of entire limbs and even pairs of
 limbs; requires precise and widespread wiring of the spinal interneurons

MOTOR SYSTEM INJURY *For coordination to be achieved among the various limbs, sets of central
Paresis- weakness, motor nerve to a muscle is damaged pattern generators *must be interconnected, *have great flexibility so that
complete paralysis -loss of motor function they can be altered quickly, *reliable methods must be available for
areflexia - motor axons cannot trigger contractions, there can be no reflexes regulating the speed of the patterns and for turning them on and off.
** Normal muscles are slightly contracted even at rest—they have some tone ** each limb has at least one central pattern generator. If one leg is
Atonia- motor nerves are transected, muscles become flaccid prevented from stepping, the other continues stepping normally
Atrophy - loss of muscle mass, absence of trophic influences from the nerves
**Complete transection of the spinal cord leads to profound paralysis below Pacemaker cells and synaptic interconnections both contribute to central
the level of the lesion pattern generation
paraplegia when only both legs are selectively affected pacemaker neurons - able to generate rhythmic activity by relying only on
hemiplegia when one side of the body is affected their intrinsic membrane conductance; membrane characteristics endow
quadriplegia when the legs, trunk, and arms are involved them with pacemaker properties that are analogous to those of cardiac muscle
s/p acute injury: cells & smooth muscle cells; primary rhythmic driving force for sets of motor
spinal shock: there is also areflexia and reduced muscle tone (hypotonia) neurons | **pacemakers are embedded w/ interconnected circuits
*temporary, after days to months is replaced by both an exaggerated muscle **the combination of intrinsic pacemaker properties and synaptic
tone (hypertonia) and heightened stretch reflexes (hyperreflexia) with interconnections that generates rhythms.
related signs—this combination is called spasticity. feedback from the muscles is not needed for the rhythms to proceed
indefinitely; rhythm generation can continue in the absence of sensory
Spinal reflexes- strongly influenced by control centers within the brain information.
Axons descend (descending control) from the brainstem & cerebral cortex &
synapse on the spinal interneurons, w/ direct input to the motor neurons. Central pattern generators in the spinal cord take advantage of sensory
descending pathways can alter the strength of reflexes feedback, interconnections among spinal segments, and interactions with
Jendrassik maneuver – done to an anxious patient, patient clasps his or her brainstem control centers
hands together and pulls; while the patient is distracted with that task, the Mammals; the rhythmic pattern is generated within the spinal cord, and
examiner tests the stretch reflexes of the leg. neurons in the brainstem control the initiation and speed of the patterns
descending pathways affects flexor reflexes sensory feedback: stretch reflex - stretching occurs on the side of the cord
mental effort – painful stimuli-tolerated, suppression of withdrawal reflexes that is currently relaxed, the effect of both stretch receptors is to terminate
anticipation - heighten the vigor of a withdrawal reflex when the stimulus activity on the contracted side of the body and to initiate contraction on the
actually arrives relaxed side
brain’s influence on spinal circuitry is thru control of spinal interneurons interconnection of spinal segments; ensures the smooth progression of
contractions down the length of the body; each segment must command its
RHYTHMIC ACTIVITY: CENTRAL PATTERN GENERATORS muscles to contract slightly later than the one anterior to it, with a lag of ~1%
Central pattern generators in the spinal cord can create a complex motor of a full activity cycle for normal forward movement
program even without sensory feedback reciprocal communication; spinal generators also inform the brainstem of
motor program - set of structured muscle commands that are determined by their activity
the nervous system before a movement begins and that can be sent to the  spinal pattern generators to produce rhythms.
muscles with the appropriate timing so that a sequence of movements occurs  sensory feedback(feedback from muscle) to modulate locomotor
without any need for sensory feedback. rhythms
central pattern generators- well studied circuits that underlie many of the  coordinate the spinal pattern generators across segments,
rhythmic motor activities that are central to animal behavior  maintain reciprocal communication between spinal generators
 and brainstem control centers. view: polka dot pattern of small dots ~0.2 mm in diameter

SPATIAL REPRESENTATIONS: SENSORY

AND MOTOR MAPS IN THE BRAIN
*spinal cord can receive sensory input, integrate it, and produce motor output Area V2/Secondary visual cortex– has series of thick and thin stripes that
that is totally independent of the brain. are separated by pale interstripes.
**neural maps- the brain organizes sensory & motor input spatially thru this **blobs and stripes seem to demarcate clusters of neurons that process and
channel different types of visual information between areas V1 and V2
The nervous system contains maps of sensory and motor information and pass them on to other visual regions of the cortex.
Sensory receptors – laid out in planar sheets, straightforward spatial
maps of the sensory environment that they encode; often project onto many Maps of somatic sensory information magnify some parts of the body
different regions of the CNS; each unique sensory surface may be mapped & more than others
remapped many times w/in the brain Homonculus – little person representing the Somatotopy(Somatosensory
*ex: sound localization, overlaying visual info w/ auditory map, mapping of the body surface of the primary somatic sensory
frequency map- hair cell in cochlea, chemical maps-do not cortex); confirmed with PET and fMRI, resembles a a trapeze artist hanging
frequency map of sound rather than a encode stimulus position upside down—the legs are hooked over the top of the postcentral gyrus and
map of the location of sounds in space dangle into the medial cortex between the hemispheres, and the trunk, upper
limbs, and head are draped over the lateral aspect of the postcentral gyrus.
The cerebral cortex has multiple visuotopic maps (1) mapping of the body surface is not always continuous
Visual field – best brain map example (2) the map is not scaled like the human body. Instead, it looks like a
Area V1 – retinotopic map; maps the visual thalamus, which in turn maps the cartoon character: the mouth, tongue, and fingers are very large,
retina, the first visuotopic map in the brain. whereas the trunk, arms, and legs are tiny.
(1)the left half of the visual field is represented on the right cortex & the *The relative size of cortex that is devoted to each body part is correlated
upper half of the visual field is represented on the lower portions of the with the density of sensory input received from that part
cortex | All the retinal axons from the left-most halves of both eyes project to *Size on the map is also related to the importance of the sensory input
the left half of the brain from that part
(2)Magnification factor scaling of the visual fields onto the visual cortex - The somatotopic maps in the cortex begin with the primary somatic sensory
is not constant | fovea(central region of visual field) - greatly magnified on axons that enter the spinal cord or the brainstem, each at the spinal segment
the cortical surface; photoreceptors& ganglion cells are densely packed @ appropriate to the site of the information that it carries. The sensory axons
the central retinal region synapse on second-order neurons, and these cells
●humans devote almost half of neocortex to visual info processing project their axons into various nuclei of the thalamus and form synapses.
●w/in Area V1 - the visuotopic maps of 2 eyes remain segregated Thalamic relay neurons in turn send their axons into the neocortex.
*in Layer IV of the primary visual cortex by having visual input derived Topographical order of body surface is maintained at each anatomical stage
from the left eye alternate every 0.25 to 0.5 mm with visual input from the
right; two sets of information, one from the left eye and one from the right The cerebral cortex has a motor map that is adjacent to and well aligned
eye, remain separated but adjacent. with the somatosensory map
Ocular dominance columns- sideview: left-right alternations look like *sensory and motor maps are adjacent and similar in basic layout; there are
columns | surface view: bands or zebra stripes myriad axonal interconnections between the primary motor and primary
**ocular dominance columns demarcate the left and right eyes somatosensory areas
Difference: neurons in the arm area of the motor cortex form distributed and
*In layer II & III - have structures called blobs, *staining for blobs: cooperative networks that control collections of arm muscles.
mitochondrial enzyme cytochrome ooxidase; sideview: round pegs | surface
superior colliculus (motor and sensory functions may even occupy the same TEMPORAL REPRESENTATIONS: TIME-MEASURING CIRCUITS
tissue) receives direct retinotopic connections from the retina as To localize sound, the brain compares the timing and intensity of input
well as input from the visual cortex to the ears
*has maps of both auditory and somatosensory information superimposed on general strategies of sound localization|mechanism by which a brainstem
its visual and motor maps circuit measures the relative timing of low-frequency sounds so that the
*motor map for orientation of the eyes is in precise register with the visual source of the sounds can be localized with precision
response map Sound localization along the vertical plane
●degree of elevation, can work well even w/ 1 ear
Sensory and motor maps are fuzzy and plastic ●depends on the shape of external ear, PINNA (where some sound reflects
Advantage of Mapping: off the curves and folds of the pinna and tragus before it enters the canal)
(1)Maps may be the most efficient way of generating nearest-neighbor ●the sound and the energy are transferred to cochlea
relationships between neurons that must be interconnected for proper fx **arcing shape of the pinna, the reflected path of sounds coming from above
*neighboring neurons most likely activated together is shorter than that of sounds from below
*synchronous activity serves to reinforce the strength of their **the direct & reflected sounds combine to create sound that are slightly
interconnections because of the inherent rules governing synaptic plasticity different upon entering the auditory canal, the combination creates
(2)Mapping may simplify establishment of the proper connections between interference pattern and spectral properties that are characteristic of the
neurons during development; easier to establish interconnections precisely elevation of the sound source.
among the neurons that represent the three sensory maps and one motor map Sound localization along the horizontal plane (azimuth)
in the superior colliculus ●requires 2 ears| Sounds must first be processed by the cochlea in each ear
(3) facilitate the effectiveness of inhibitory connections; construct an edge- then compared by neurons w/in the CNS to estimate horizontal direction
detector circuit| (edge detection) is heightened by lateral connections that What is compared?
suppress the activity of neurons representing the space slightly away from the High Frequency Sounds Low frequency sounds
edge ●interaural intensity difference ●measures interaural delay
Disadvantage of Mapping: (ear to ear)|the ear facing the sound ●Sounds below ~2 kHz have a
(1)Maps are abstract representations; distorted by the shortcomings of hears it as louder than the ear facing wavelength that islonger than the
experimental measurements away because the head casts a width of the head itself.
(2) Maps of sensory space onto a brain area are not point-to-point “sound shadow” *no occurrence of interaural
representations; 1 point in sensory space-activates a large group of neuron; *straight sound – same intensity intentisty
strength of activation is most intense within the center of the activated (sounds coming from infront and
neuronal group, but the population of more weakly activated neurons may behind may have similar intensity) *front-back ambiguity
encompass a large portion of an entire brain.
(3)maps may change with time- dynamic and can be reorganized rapidly and
substantially as a function of development, behavioral state, training, or
damage to the brain or periphery (PLASTICITY)
In damage: severed nerve to become remapped to another body part.
 *each is tuned to a different interaural delay and a different sound locale
along the horizontal axis
The brain measures interaural timing by a combination of neural delay
lines and coincidence detectors Arrangement of delay lines  orderly spatial mapping of sound direction
Detection of intraural timing(very small time differences) – via precise delay lines, synaptic inhibition, and perhaps other neuronal properties
arrangement of neurons in space. combine to optimize the measurement of timing in mammals.
Cochlea – no map for sound location, CNS localizes low frequency sounds by
(each) Cochlear Nuclei neuron medial superior olivary (MSO) calculating an interaural time-delay map, using information
nucleus neuron from both ears together
receive information from only the ear receive abundant input from both
on that one side ears, compare the timing (the phase) Inferior colliculus receives parallel information on both timing
of sounds arriving at the two ears delay and intensity difference; it transforms these two sets of
cochlear nucleus neurons are exquisitely sensitive to interaural information, combines them, and produces a complete map
activated by auditory stimuli, their time delay, and the optimal delay for of sound direction.
action potentials tend to fire with a superior olivary neurons varies
particular phase relationship to the systematically across the nucleus
sound stimulus.
its firing is phase locked to the sound Has a spatial map of interaural delay,
waves, at least for relatively low systematic map of sound frequency,
frequencies. so it simultaneously maps two
qualities of sound stimuli.
Cochlear neurons preserve
the timing information of sound
stimuli

Medial Superior Olivary neurons

tuning of MSO neurons to interaural delay seems to depend on neural
circuitry that combines “delay lines” with “coincidence detection”
Delay lines are the axons from each cochlear nucleus; their length and
conduction velocity determine how long it takes sound-activated action
potentials to go from a cochlear nucleus to the axon’s presynaptic terminals
onto MSO neurons | The difference in conduction delay between the axon
from the right side and that from the left side determines the optimal
interaural delay for that particular olivary neuron.
coincidence detector- is the olivary neuron; only when action potentials
from both the left- and right-ear axons reach the postsynaptic MSO neuron
simultaneously is that neuron likely to receive enough excitatory synaptic
transmitter to trigger an action potential.
Will fire if: coincidence between input from the left and right
Neuron will not fire If: input from the two ears arrives at the neuron out of
phase, without coincidence in time
*Neurons display coincidence for different interaural delays.