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Interpretive Framework: GLA — Downstream Amplifier Module
Shear-Activated PEM Generation Module

Shear-Activated PEM Generation Module

(Endothelial Timing Failure → Skeletal Muscle Injury → Delayed PEM)

Author: Michael Daniels· Framework: GLA·2.5 · Date: January 14th 2026 · This document presents a systems-level interpretation and is not clinical guidance or a treatment recommendation.

Snapshot

Module purpose

This module defines the core mechanism by which post-exertional malaise (PEM) is generated in ME/CFS: a shear-activated failure state arising from impaired endothelial signal-timing precision that produces skeletal-muscle oxygen-extraction failure during exertion and delayed cellular injury during recovery.

Scope

Interpretive guardrail

This module addresses how PEM is generated once a vulnerable control state exists; it does not propose shear stress, exertion, or skeletal muscle as the initiating cause of ME/CFS.

Orientation Note — How to Read This Module
This module should be read as a mechanistic explanation of how post-exertional malaise (PEM) is generated once a vulnerable physiological control state exists, not as a claim about the original cause of ME/CFS.

Key points for interpretation:
Exertion and shear stress are activators, not causes.
The processes described here explain how normal physiological demands trigger PEM in a system with impaired control precision. They do not imply that exertion, fitness level, or skeletal muscle pathology initiates the disease.

The focus is timing and distribution, not capacity.
This module addresses failures of signal timing, flow distribution, and recovery coordination — not global deficits in oxygen delivery, cardiac output, lung function, or mitochondrial quantity.

Skeletal muscle is treated as the primary PEM generator, not the primary disease site.
Muscle is highlighted because it is uniquely sensitive to microvascular timing errors and capable of encoding delayed injury. This does not exclude parallel involvement of immune, autonomic, or central nervous system processes.

Recovery is the critical injury window.
Although the triggering event occurs during exertion, the model explicitly places the major tissue injury and symptom amplification during the recovery phase, explaining the delayed onset of PEM.

This module integrates with, but does not replace, other GLA components.
Immune activation, autonomic dysregulation, metabolic stress, membrane instability (e.g., SMPDL3B), buffering capacity (e.g., haptoglobin), and disease phase all condition susceptibility and severity. This module explains how those upstream factors are converted into the defining clinical feature of PEM.

In short: this module explains why PEM is delayed, disproportionate, and reproducible, even when standard cardiopulmonary measures appear normal.
Module placement within GLA
Primary layer coupling
• Layer 2 — Membrane / Execution Surface Stability
SMPDL3B-mediated lipid microdomain integrity and caveolar anchoring
• Layer 4 — Distribution & Buffering
Caveolin-1–dependent signal timing, nitric oxide spatial precision, and vascular/RBC buffering capacity (e.g., haptoglobin)
• Layer 5 — Recovery Bandwidth
ER–mitochondrial calcium handling, oxidative stress resolution, and post-exertional recovery capacity

Functional role within the GLA system
• Primary trigger organ: Skeletal muscle
• Primary physiological activator: Normal physiological shear stress
• Primary failure mode: Loss of timing precision and signal coordination (not loss of delivery capacity, energy supply, or organ reserve)
Upstream context — where this module sits in the GLA hierarchy
The Shear-Activated PEM Generation Module operates downstream of earlier GLA control layers. These upstream layers establish the vulnerable physiological state but do not, on their own, generate post-exertional malaise.

Layer 0 — Genetic & baseline control bias
Polygenic modifiers shape immune signal gain, membrane maintenance, lipid routing, vascular buffering, and recovery bandwidth. These influences bias susceptibility and failure mode (e.g., control fragility vs overshoot) but are not symptomatic by themselves.

Layer 1 — Immune control & signal termination
Impaired resolution of innate immune signaling (e.g., prolonged priming, poor termination, ER stress coupling) establishes a persistent “activated but non-inflammatory” control state. This layer conditions downstream membranes and endothelial timing without directly causing PEM.

Transition to execution
When Layer 1 control instability persists, downstream execution surfaces (Layers 2–5) become increasingly vulnerable to normal physiological stressors. The present module explains how that vulnerability is converted into delayed skeletal-muscle injury and PEM.
Execution-layer overview — Layers 2–5 at a glance
Layer 2 — Membrane & execution surface stability
Governs the physical integrity and signaling reliability of lipid microdomains (e.g., SMPDL3B-dependent raft anchoring). Instability here lowers tolerance to mechanical and biochemical stress without creating symptoms on its own.

Layer 3 — Metabolic routing & substrate handling
Shapes how energy substrates, lipids, and metabolites are routed and buffered during demand and recovery. Constraints at this layer limit flexibility under stress but do not directly determine where injury occurs.

Layer 4 — Distribution, timing & buffering
Controls endothelial signal timing, nitric oxide spatial precision, microvascular flow topology, and vascular/RBC buffering. Failure here converts normal shear stress into heterogeneous perfusion and extraction mismatch.

Layer 5 — Recovery bandwidth & reset capacity
Determines whether tissues can resolve post-exertional stress through effective calcium handling, redox control, and mitochondrial recovery. When constrained, injury is encoded during recovery rather than during exertion itself.
Canonical placement statement
This module occupies the interface between endothelial control timing (Layers 2–4) and post-exertional recovery failure (Layer 5), explaining how upstream membrane and distribution instability is converted into delayed skeletal-muscle injury and PEM.

Section 1 — Inputs: Physiological Shear Stress

Physiological shear stress rises normally in response to everyday demands, including:

These inputs are continuous features of normal human physiology and are present even at low levels of activity.

Key framing
Shear stress is a normal physiological input and does not represent excessive exertion, pathological overload, or abnormal mechanical force.

Implications

This framing explicitly excludes post-exertional malaise (PEM) as a consequence of:

Instead, shear stress functions in this module as a physiological activator that reveals an underlying failure of endothelial signal decoding and timing precision.

Section 2 — Control Surface: SMPDL3B → Cav-1 → eNOS

Core control architecture

Endothelial shear sensing and response depend on a precisely organized membrane control surface. This control surface translates mechanical shear into coordinated biochemical signals through tightly regulated spatial and temporal organization.

Together, this system translates mechanical shear into precisely timed, spatially restricted endothelial responses, rather than continuous or diffuse signaling.

Key clarification
Nitric oxide signaling failure in ME/CFS reflects loss of timing precision and spatial localization, not global nitric oxide depletion or impaired production capacity.

Failure mode integration

In ME/CFS-relevant control states, this membrane-based timing architecture becomes unstable:

This loss of timing control at the membrane–caveolar interface constitutes the root control failure for this module.

Figure 1. Endothelial control surface timing architecture (SMPDL3B → Cav-1 → eNOS)

Shear is decoded into precisely timed, spatially localized responses—not diffuse NO “power.”

GLA · Layers 2–4
Control-surface timing: decode shear → precise endothelial response Timing • localization • termination (not global NO depletion or “more NO”) Physiological shear input standing • exertion • heat • autonomic drive SMPDL3B stabilizes lipid microdomains & caveolar integrity Cav-1 brake / release gate eNOS localized NO pulses Timing brake / release coordination Localization restricted microdomain signaling Termination signal shutoff & reset Key claim: NO signaling failure reflects loss of timing precision & spatial localization — not global NO depletion.

Caption. Shear sensing depends on a membrane control surface in which SMPDL3B stabilizes lipid microdomains and caveolar structure, Cav-1 gates timing (brake/release), and eNOS generates localized nitric oxide pulses. In this module, dysfunction reflects loss of timing and localization rather than reduced nitric oxide production capacity.

Section 3 — Endothelial Output: Flow Topology Failure

Mistimed endothelial signaling produces a characteristic failure of microvascular flow topology, rather than a reduction in total blood flow.

As a consequence of impaired Cav-1–eNOS timing, capillary-level flow heterogeneity develops, characterized by:

This pattern reflects a loss of coordinated capillary recruitment and flow smoothing at the microvascular level.

Key clarification
The endothelial failure is topological, not scalar: overall blood flow may increase appropriately, but it is distributed incorrectly across the microvasculature.

What this mechanism does not involve

This flow-topology failure does not involve:

Instead, the defining abnormality is misallocation of flow in space and time, setting the conditions for downstream oxygen-extraction failure despite preserved central delivery.

Figure 2. Microvascular flow topology failure despite preserved total flow

Overall inflow can be appropriate while distribution across the microvasculature is incorrect.

Flow topology
Topological vs scalar endothelial failure Same total inflow → different microvascular distribution Normal capillary flow Fast channel (high velocity) Silent / under-perfused region Panel A Normal distribution Panel B Flow topology failure Inflow (same) Inflow (same) Even recruitment • smooth velocities • consistent transit times Fast channels + silent regions • shortened transit in over-perfused paths • unstable delivery elsewhere Total flow preserved — distribution failed (topological failure, not scalar hypoperfusion)

Caption. Microvascular dysfunction in this module is defined by flow topology failure: total inflow can remain appropriate while perfusion becomes spatially heterogeneous. Panel A illustrates even recruitment and smooth velocities. Panel B illustrates over-perfused fast channels alongside under-perfused or intermittently silent regions. This topological misallocation sets the stage for tissue-level oxygen extraction failure despite preserved central delivery.

Section 4 — Why Skeletal Muscle Is the Trigger Site

Skeletal muscle is uniquely vulnerable to endothelial timing and distribution failures because it combines extreme perfusion demands with strict spatial precision requirements.

Specifically, skeletal muscle:

As a result, even modest microvascular timing errors can produce disproportionate physiological consequences in contracting skeletal muscle.

Key clarification
The same endothelial timing error tolerated by other organs becomes pathological in contracting skeletal muscle.

Functional implication

This vulnerability positions skeletal muscle as the primary generator of post-exertional malaise (PEM), where endothelial control failure is first converted into tissue-level stress and delayed injury — rather than as a passive downstream recipient of systemic dysfunction.

Table 1. Relative dynamic perfusion range across major organ systems

Skeletal muscle has the largest dynamic perfusion range. This makes it the most sensitive tissue for exposing endothelial timing and distribution failures during activity. Values are approximate and depend on intensity, posture, thermoregulation, and training status.

Organ / vascular bed Typical change during exercise Why it matters for PEM framing
Skeletal muscle ~20–50× (rest → maximal, per active muscle) Largest demand swing → most sensitive to timing/distribution errors and capillary recruitment failure
Coronary (heart) ~3–5× Tight autoregulation → smaller dynamic range, less likely to expose topology failure at low exertion
Brain ~0.5–1.5× Relatively stable flow; small distribution errors can still cause symptoms but demand swing is modest
Skin ~0–10× (heat-dependent) Thermoregulatory variability; can amplify symptoms but is not the dominant extraction bed
Splanchnic (GI/liver) (often decreases) Redistribution bed; contributes to post-exertional strain but not the primary site of extraction failure
Kidney (often decreases) Perfusion trade-off impacts volume regulation and recovery bandwidth rather than exertion-time extraction

Interpretation. PEM is most reliably generated where perfusion demand swings are largest and capillary recruitment must be precisely coordinated. Skeletal muscle uniquely meets both conditions, which is why the same endothelial timing error tolerated elsewhere becomes pathological in contracting muscle.

Section 5 — Exertion Phase: Oxygen Extraction Failure

During exertion, systemic oxygen delivery functions appropriately, while local utilization in skeletal muscle fails.

Central delivery (preserved)

These findings confirm that oxygen is successfully loaded, transported, and delivered at the systemic level.

Microvascular execution (failed)

Despite preserved central delivery:

The result is a failure of localized oxygen extraction at the tissue level.

Physiological consequence

Evidence alignment

Key summary
Oxygen delivery occurs; oxygen use fails.

Figure 3. Preserved oxygen delivery with failed skeletal-muscle extraction

Central delivery rises appropriately, but microvascular distribution and unloading fail at the muscle level.

iCPET pattern
Exertion-time physiology: delivery preserved, extraction fails The iCPET signature: high venous O₂ during effort despite rising cardiac output Central delivery (preserved) Microvascular extraction (failed) Cardiac output increases appropriately with workload Arterial O₂ remains normal (gas exchange intact) normal Systemic transport O₂ is loaded, carried, delivered Skeletal muscle capillary bed distribution heterogeneity → shortened transit in fast channels → poor unloading Transit time ↓ in over-perfused channels Arterial O₂: delivered to muscle Mixed venous O₂: remains high during effort poor unloading Delivery intact — utilization failed (peripheral extraction failure)

Caption. During exertion, central oxygen delivery can remain appropriate: cardiac output rises with workload and arterial oxygenation remains normal. However, microvascular distribution in skeletal muscle becomes heterogeneous, shortening red blood cell transit time in over-perfused channels and limiting oxygen unloading. The characteristic iCPET pattern emerges: elevated mixed venous oxygen saturation during effort, indicating preserved delivery but impaired peripheral extraction.

Section 6 — Latent Injury Encoding (Exertion, Not Yet PEM)

During exertion, skeletal muscle does not immediately fail, despite impaired oxygen extraction. Instead, it transitions into a compensated but unstable physiological state that temporarily sustains function while encoding latent injury.

This compensated state is characterized by:

These compensatory responses preserve short-term performance but increase cellular stress and reduce recovery headroom.

Key clarification
Exertion reveals a pre-existing failure state but does not cause immediate tissue injury.

Functional consequences

This latent, compensated state explains:

The critical transition from compensation to injury occurs after exertion ends, during the recovery phase addressed in Section 7.

Figure 4. Compensated exertion state and latent injury encoding

During exertion, function can be temporarily maintained while cellular stress is encoded for recovery-phase injury.

Latency
Latent injury encoding: compensated exertion state False OK window: performance may persist while recovery headroom is reduced Exertion window compensation holds function Early recovery transition toward injury amplification Compensated state during exertion Intracellular Ca ²⁺ influx increases to maintain contraction Glycolysis upregulated to compensate for inefficient oxidative use Mitochondrial workload increases despite impaired oxygen utilization Latent encoding recovery headroom shrinks ER–mitochondrial coupling tightens raises vulnerability to stress during recovery Latent injury encoding cellular stress increases, but injury is not yet symptomatic “False OK” window Function can persist while recovery headroom shrinks — the critical transition occurs after exertion ends.

Caption. During exertion, skeletal muscle can enter a compensated but unstable state that maintains function despite impaired oxygen extraction. Compensation is supported by increased intracellular Ca2+ influx, increased glycolysis, and increased mitochondrial workload. At the same time, ER–mitochondrial coupling tightens and recovery headroom shrinks, encoding latent injury that may not be symptomatic during exertion. This explains the “false OK” window and sets up the recovery-phase amplification described in Section 7.

Section 7 — Recovery Phase: Injury Amplification

After exertion ends, systemic demand for perfusion decreases, but the upstream endothelial control failure does not immediately resolve.

During early recovery:

This mismatch converts the previously compensated stress state into active injury.

Downstream consequences

The persistence of flow heterogeneity during recovery produces:

These processes drive progressive tissue injury after activity has stopped, rather than during exertion itself.

Key summary
Recovery, not exertion, is where collapse occurs.

Figure 5. Recovery-phase amplification of microvascular and cellular injury

After activity stops, demand falls—but timing/topology failures can persist, converting compensated stress into injury.

Recovery collapse
Recovery-phase amplification: mismatch persists after demand falls The compensated exertion state transitions into injury during recovery Perfusion demand falls contractile activity stops → systemic demand decreases Flow heterogeneity persists timing/topology failure does not immediately resolve → mismatch remains persistent mismatch converts stress → injury Micro-ischemia / reperfusion mismatch under-perfused regions receive delayed or excessive flow ROS bursts oxidative stress amplification ER–mitochondrial Ca ²⁺ dysregulation prevents reset of metabolism and signaling Repair and resolution fail to complete impaired cellular repair and resolution signaling → recovery bandwidth is limited → injury accumulates after activity ends repair Recovery, not exertion, is where collapse occurs

Caption. After exertion ends, perfusion demand decreases, but the upstream endothelial timing failure does not necessarily resolve immediately. Microvascular flow topology can remain heterogeneous, maintaining a mismatch between flow and tissue needs. This persistence converts the compensated exertion state into active injury via micro-ischemia/reperfusion mismatch, reactive oxygen species (ROS) bursts, sustained ER–mitochondrial Ca2+ dysregulation, and impaired repair and resolution signaling. The result is progressive tissue injury during recovery, not during exertion itself.

Section 8 — Clinical Expression: Post-Exertional Malaise

Over the ensuing hours to days following exertion and recovery-phase injury amplification, structural and metabolic injury accumulates in skeletal muscle, reflecting unresolved cellular stress and impaired repair.

Whole-body symptoms emerge as post-exertional malaise (PEM), including fatigue, pain, cognitive dysfunction, autonomic instability, and reduced functional capacity.

The delayed onset and duration of PEM reflect the time course of tissue injury, stress signaling, and incomplete recovery, rather than acute exertional failure.

Determinants of severity and duration

The intensity and persistence of PEM are shaped by:

Key clarification
Post-exertional malaise reflects delayed tissue injury following a shear-activated control failure, not immediate energy depletion or acute exertional exhaustion.

This framework does not claim that PEM is caused by exercise itself, does not assert a universal primary muscle disease, does not depend on nitric oxide deficiency, and does not exclude contributions from immune or autonomic dysfunction.

Figure 6. Shear-activated PEM generation timeline

One-panel synthesis: how normal shear input becomes delayed, multi-system PEM via timing failure and recovery-phase injury amplification.

Synthesis
Shear-activated PEM generation: integrated timeline Exertion reveals the failure state; recovery amplifies injury; PEM emerges with delay Exertion phase Recovery phase Symptom phase Shear input Timing failure Topology failure Extraction failure Latent injury Physiological activator Control-surface decoding fails Distribution misallocated iCPET: venous O₂ high “False OK” window Recovery-phase collapse ischemia/reperfusion mismatch • ROS bursts • ER–mitochondrial Ca²⁺ dysregulation • repair failure Post-exertional malaise (PEM) delayed fatigue • pain • cognitive dysfunction autonomic instability • reduced capacity Exertion reveals; recovery amplifies; PEM emerges with delay

Caption. Integrated synthesis of the module logic. Physiological shear input is decoded by an endothelial control surface that requires timing precision. When timing fails, microvascular flow becomes topologically misallocated, producing skeletal-muscle oxygen extraction failure during exertion (iCPET pattern). Muscle can remain temporarily functional in a compensated state while latent injury is encoded. After exertion ends, persistent flow heterogeneity during recovery drives ischemia/reperfusion mismatch, ROS bursts, sustained ER–mitochondrial Ca2+ dysregulation, and impaired repair—leading to delayed, multi-system post-exertional malaise (PEM).

Table 2. Determinants of PEM severity and duration

PEM severity reflects how much injury is amplified during recovery and how long repair and resolution remain incomplete. The same exertional input can produce very different outcomes depending on baseline control state, buffering capacity, and accumulated prior injury.

Determinant What it represents in this module Effect on PEM severity / duration Typical pattern
Disease phase (baseline control state) Baseline stability of timing, distribution, and recovery coordination — i.e., how close the system is to a failure threshold before shear input. Lower baseline control increases likelihood that normal shear triggers topology failure, and lengthens the recovery window required for repair and reset. Early-phase: intermittent tolerance with clearer recovery.
Late-phase: low threshold, prolonged or continuous instability.
Physiological buffering capacity (e.g., haptoglobin) Ability to absorb downstream consequences of shear-related stress (vascular/RBC buffering, hemoglobin/heme handling, oxidative load containment). Lower buffering increases oxidative amplification and slows resolution, raising symptom intensity and prolonging PEM duration under the same stressor. Better buffering: higher tolerance and faster recovery.
Weaker buffering: sharper crashes and longer recovery tails.
Prior injury burden (cumulative baseline erosion) Pre-existing structural/metabolic strain and reduced repair bandwidth from previous episodes — a reduced “recovery reserve.” Higher prior injury lowers the threshold for subsequent PEM and increases the probability of stepwise worsening, with longer and deeper post-exertional impairment. Low burden: clearer “baseline return.”
High burden: incomplete return and progressive threshold erosion.

Interpretation. PEM severity is not determined only by the size of the exertional input. It is determined by whether shear variability exceeds shear tolerance, how strongly recovery-phase injury is amplified, and how much recovery bandwidth remains to complete repair and reset.

Interpretive Framework Documents (GLA v2.1 → v2.5)

The documents listed below define the conceptual and methodological framework used to interpret genetic signals and physiological mechanisms in this paper. Collectively, they establish layer boundaries, phenotype discipline, and phase dependence within the Gut–Liver–Autonomic (GLA) system architecture.

These materials are provided for transparency and interpretive context only. They are not cited as evidentiary sources and should be read as evolving systems-biology models used to organize and constrain interpretation, rather than as claims of mechanism or causation.

Core GLA framework documents

SMPDL3B phenotype frameworks

Control layers & system modulators