SAQPhysiologyNeurophysiology2001 · Pain pathway and primary hyperalgesia

Question bank · 2001 · Physiology

The pathway explains why an injured finger hurts.
Primary hyperalgesia explains why it goes on hurting at a touch.

Show the model answerAttempt it first — that is what makes it stick

The pathway from the finger

What earns the marks

ReceptorFree nerve endings; transduction to action potentials
FibresAδ first pain, C second pain, in the digital nerves
First-order neuroneCell body in the dorsal root ganglion, C6 to C8; synapse in the dorsal horn
Second-order neuroneCrosses; contralateral spinothalamic tract to the thalamus
Third-order neuroneThalamus to somatosensory cortex

Read the question: Briefly outline” is a limit. One line per neurone and a labelled diagram answer it; the remaining two sentences of the question, a definition and a mechanism, are where the detail belongs.

Pathway

The ascending pathway, in three neurons

Read it from the bottom. The first-order neuron has its cell body in the dorsal root ganglion, outside the cord, and synapses in the dorsal horn. The second-order neuron starts in the dorsal horn, crosses in the cord and ascends on the opposite side. At the brainstem the projection divides: the lateral, neospinothalamic route relays in the ventral posterolateral nucleus (VPL) and reaches the primary somatosensory cortex, which is where and how intense; the medial, paleospinothalamic route passes through the reticular formation and periaqueductal grey to the intralaminar nuclei and on to the cingulate and insula, which is how unpleasant. The third-order neuron runs from thalamus to cortex. From the face the plan is the same, with the first-order cell body in the trigeminal ganglion, the second in the spinal trigeminal nucleus and the third in the ventral posteromedial nucleus.

CortexS1, S2, insula, anterior cingulateThalamusVPL (lateral) and intralaminar (medial)BrainstemReticular formation, periaqueductal greySpinal cordDorsal horn, laminae I, II and VPeripheryFree nerve endings of nociceptorsmidlineDorsal root ganglionfirst-order cell bodyDorsal horncrosses hereReticular formationand PAGVPLIntralaminarS1: locationCingulate, insula: affect123NeospinothalamicPaleospinothalamic
StepStructureDetail
TransductionNociceptors in the skin of the fingerFree nerve endings. Noxious heat, pressure or chemicals open TRPV1, ASIC and P2X3 channels; the generator potential fires action potentials through voltage-gated Na⁺ channels
FibresDigital nerves, branches of the median, ulnar and radial nervesAδ, thinly myelinated, 12 to 30 m/s: sharp, localised first pain. C, unmyelinated, 0.5 to 2 m/s: dull, burning second pain
First-order neuroneCell body in the dorsal root ganglion: C6 for the thumb, C7 for the middle three fingers, C8 for the little fingerEnters the cervical cord by the dorsal root, runs 2 to 3 segments in Lissauer's tract, and synapses in the dorsal horn: Aδ in laminae I and V, C in lamina II. Glutamate and substance P
Second-order neuroneCell body in the dorsal hornCrosses in the ventral white commissure within one or two segments, and ascends in the contralateral anterolateral quadrant as the spinothalamic tract
Third-order neuroneCell body in the thalamusVentral posterolateral nucleus, through the internal capsule, to the primary and secondary somatosensory cortex of the opposite hemisphere: where and how intense. Medial projections to the cingulate and insula: how unpleasant

In the dorsal horn, Aδ input also initiates the rapid withdrawal of the finger through a spinal reflex, and the difference in conduction velocity is why a sharp pain is felt first and a dull ache follows it.

What is primary hyperalgesia?

What earns the marks

HyperalgesiaIncreased pain from a stimulus that normally hurts: a leftward shift of the stimulus-response curve
PrimaryAt the site of injury, from sensitisation of the nociceptors there
FeaturesLower threshold, more pain above threshold, spontaneous pain; to heat and to mechanical stimuli
The contrastSecondary hyperalgesia: uninjured tissue around the injury, central, mechanical only

Read the question: “What is” asks for the definition and what marks it out. Primary hyperalgesia is peripheral and extends to heat; secondary hyperalgesia is central and mechanical only. Without that contrast, “primary” has not been explained.

Primary hyperalgesiaSecondary hyperalgesia
WhereThe injured tissueThe uninjured tissue around it
Where the change isThe nociceptor terminal: peripheral sensitisationThe dorsal horn: central sensitisation
More pain fromHeat and mechanical stimuliMechanical stimuli only
How it showsLower pain threshold; more pain from stimuli above threshold; spontaneous painAllodynia and tenderness to touch
Sensitisation

Two zones around an injury, and what sensitisation does to the response

A. Primary hyperalgesia is in the injured tissue and is peripheral; secondary hyperalgesia is in the uninjured tissue around it and is central. The discriminator is heat: the primary zone is more sensitive to thermal and mechanical stimuli, the secondary zone to mechanical stimuli only. B. Sensitisation moves the stimulus-response curve left and up. Left is a lower threshold, so a stimulus that was not painful now is (allodynia); up is more pain from a stimulus that already was (hyperalgesia). The axes are deliberately unnumbered: the sources give the direction of the shift, not a scale.

A · Where each zone isInjuryPrimary hyperalgesiainjured tissue · peripheralSecondary hyperalgesiauninjured tissue · centralB · What it does to the responsePain intensityStimulus intensitynormal thresholdnew thresholdAllodyniaHyperalgesiasame stimulus, more pain

The mechanisms by which primary hyperalgesia develops

What earns the marks

The mediatorsReleased from damaged cells, mast cells, platelets and immune cells, or made at the site
Activation against sensitisationSome make the terminal fire; others lower its threshold
The cellular mechanismG-protein-coupled receptors, second messengers and kinases phosphorylate TRPV1 and Na⁺ channels
What changes in the nociceptorLower threshold, greater discharge, spontaneous discharge
RecruitmentSilent nociceptors become responsive
SpreadNeurogenic inflammation: substance P, CGRP and neurokinin A released from the terminals
Gene expressionNerve growth factor, through trkA, increases synthesis of neuropeptides and ion channels

Read the question: The mechanism asked for is peripheral. Wind-up, the NMDA receptor and central sensitisation explain secondary hyperalgesia; here they are worth a sentence of contrast at most.

How injury sensitises the nociceptor

Injury
Cells are damaged; platelets, mast cells and immune cells are activated.

The inflammatory soup

Released
K⁺, H⁺ and ATP from damaged cells; histamine from mast cells; serotonin from platelets.
Made at the site
Bradykinin; prostaglandins, with cyclo-oxygenase upregulated by cytokines and growth factors; leukotrienes; cytokines; nerve growth factor.

Acting on the terminal

Directly on ion channels
Protons and serotonin open channels such as ASIC and 5-HT3: the terminal is activated.
Through receptors
Most bind G-protein-coupled receptors (bradykinin B2, prostanoid EP). Second messengers activate kinases that phosphorylate TRPV1 and Na⁺ channels.
The nociceptor is sensitised
Its threshold falls, it discharges faster when stimulated, and it discharges spontaneously. Silent nociceptors become responsive.
The response spreads
Impulses run antidromically into the fibre's other branches and release substance P, CGRP and neurokinin A: vasodilatation, plasma extravasation, more bradykinin, histamine from mast cells and serotonin from platelets.
Primary hyperalgesia
At the injured site, low-intensity mechanical stimuli now hurt, heat hurts more, and stimuli that already hurt hurt more.
The mediators act synergistically, and most are not stored but made at the site once injury has occurred.

Activation and sensitisation are different actions

Activate: make the terminal fireSensitise: lower its threshold
PotassiumProstaglandins
SerotoninLeukotrienes
BradykininSubstance P
Hydrogen ionNoradrenaline
HistamineNeurokinin A and B
ATP and adenosineCalcitonin gene-related peptide
Nitric oxide
  • Bradykinin does both, through its B2 receptor.
  • Prostaglandins, mainly PGE2, sensitise rather than stimulate: through EP receptors they phosphorylate Na⁺ channels and TRPV1, so the terminal responds to mechanical, chemical and thermal stimuli it previously ignored. This is why an NSAID acts on the hyperalgesia of inflammation rather than on the stimulus itself.

Mechanisms that sustain it

MechanismHowEffect
Channels already present are modifiedPhosphorylation through second messengers and kinasesLower threshold and greater discharge of the nociceptors already active
More afferents are recruitedSilent nociceptors, unresponsive in normal tissue, respond once it is inflamedMore fibres firing, as well as each fibre firing more
Mediators amplify one anotherCytokines from immune cells release other mediators: interleukin-1β and interleukin-6 release prostaglandins; tumour necrosis factor α and interleukin-1β stimulate nerve growth factor productionThe soup keeps itself going while inflammation lasts
Gene expression changesNerve growth factor, from fibroblasts, Schwann cells and keratinocytes, acts through trkA and alters gene transcriptionMore neuropeptides (neurokinins and CGRP) and ion channels are made. Receptors and channels on the nociceptor, TRPV1 and Nav1.8 among them, adapt in ways that lower its firing threshold
ResolutionInflammation settles as the tissue healsPeripheral sensitisation diminishes and nociceptors return to their resting threshold

Primary hyperalgesia is protective: it helps prevent tissue that is already damaged from being damaged further. It becomes a problem when the inflammation does not resolve and the sensitised periphery keeps driving the dorsal horn.

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