REF-1355 · Magnusson & Kjaer, 2019 · The Journal of Physiology · Paper · PubMed · Free full text

Why does my Achilles lag behind my fitness when I start running?

Its core barely renews after adolescence; it adapts to high strain more than moderate, and loses stiffness in 1-2 weeks of limb unloading. Ramp load slowly (practice).

Main caveat: Narrative review. No search method or appraisal; statements are the authors' synthesis, and many draw on the Copenhagen group's own studies.
moderate evidenceMeasured on non-athletes to trained athletes

Magnusson and Kjaer review how human tendon responds to loading, unloading, ageing and injury.

On turnover, they set two lines of evidence side by side: stable-isotope biopsy studies found collagen synthesis of about 1% per 24 h in one study and around 0.2% per 24 h in others, while bomb-pulse 14C dating showed that the core of the human Achilles tendon changes until about 17 years of age and is stable thereafter (REF-1356).

They reconcile them with a suggested two-pool model: a large stable pool formed in childhood and adolescence, and a much smaller pool with fast turnover for daily maintenance, whose location is unknown.

On loading, the cellular response is well documented in rodents and cell cultures, but in adult human tendon an acute rise in collagen synthesis seen in one study could not be robustly confirmed by several others; the authors suggest adult human tendon is far less responsive than that of growing rodents and that new tissue may be laid down at the periphery "analogous to growth of a tree".

Tendon does hypertrophy with long-term loading — the most convincing human evidence is a 20% larger tendon cross-section on the side of athletes who had a 22% side-to-side strength difference — and in healthy Achilles tendons loading at 90% of maximal voluntary contraction (~5% strain) raised stiffness and cross-section more than 55% (~3% strain) did; contraction mode (concentric or eccentric) did not matter.

Unloading works fast: 2-3 weeks of immobilisation cut the collagen synthesis rate by 80%, and 1-2 weeks of limb unloading lowered tendon stiffness with no change in its size. With age the tendon is not lost, and a lower modulus in older people disappeared when they were matched with young people for activity.

A surgically repaired Achilles tendon still showed raised glucose uptake 12 months after repair, and the authors conclude that the tendon's response to loading after injury is rather slow, not normalised until sometime after 6-12 months.

Our reading: this is the review behind "tendon is the slow tissue" — slow to renew and slow to heal — with an asymmetry worth keeping: stiffness is lost within one or two weeks of limb unloading or immobilisation, while the evidence for gains comes from training studies of several months; it does not give a time course for stiffness gains, and the authors note that studies of recovery between loading sessions are lacking.

Protocol

Design: narrative topical review; no search method, inclusion criteria or pooling; n = not applicable (the number of studies and participants behind each statement is not given).
Scope: cellular and molecular, then structural and mechanical, responses of tendon to loading, unloading, ageing and injury (tendinopathy and rupture), with a summary table of the overall direction of each response.
Evidence mix: cell cultures and engineered tendon constructs; rodents (often still growing, 10-12 weeks old in the loading studies), horses and rabbits; human microdialysis, stable-isotope biopsy, bomb-pulse, imaging and in vivo mechanical studies.
Funding and interests: "supported by the Danish Council for Independent Research, Lundbeck Foundation and the Center for Healthy Aging"; competing interests "None declared". Crossref's funder metadata adds the Nordea Foundation (NF-007) with the Lundbeck grant (R198-2015-207), and Europe PMC lists the Novo Nordisk Foundation (NNF16OC0022846); none is a device maker, federation or data platform.

Key Results

QuestionWhat the review statesWhat it rests on
Collagen synthesis in human tendon~1% per 24 h in one study; around 0.2% per 24 h in othersstable-isotope infusion with tendon biopsy
Lifetime turnover of the coreAchilles core collagen changes until about 17 years, then stable; horse tendon half-life about 200 years (high-strain) vs ~34 years (low-strain)14C bomb pulse (REF-1356); racemization in horses
Acute loading → collagen synthesis in adult tendonraised in one study; not robustly confirmed by several others using the same method and exercise; mid-tendon gene expression fell or did not changehuman biopsy studies
Long-term loading → hypertrophya 22% side-to-side strength difference went with a 20% larger tendon cross-section on the stronger side ("perhaps currently some of the most convincing evidence")athletes loading one side more for years
Strain dose90% MVC (~5% strain) raised stiffness and cross-section vs 55% MVC (~3% strain); long-duration loading (6 s cycle) more than faster 2 s cyclestraining studies in healthy Achilles tendons
Contraction mode12 weeks of concentric or eccentric knee extension gave similar tendon hypertrophy; Achilles stretch is identical in a heel rise and a heel drophuman training and imaging studies
Unloading2-3 weeks of immobilisation: 80% lower collagen synthesis rate; 1-2 weeks of unloading: lower stiffness, no change in cross-sectionhuman immobilisation studies
Ageingno loss of tendon tissue with age; a lower modulus in old (66 y) than young (26 y) people disappeared when matched for activitycross-sectional human studies
After rupture repairglucose uptake 6x at 3 months, 3x at 6 months, 1.6x at 12 months that of intact tendon; normalised "sometime after 6–12 months"human imaging study
Recovery between loading sessions"Studies specifically addressing the effects of recovery and how it affects tendon adaptation are lacking"—

Summary table of the review (arrows = overall direction; anabolic/catabolic · cell number · tissue synthesis · cross-section · stiffness): loading ↑ · → · ↑ · ↑ · ↑; unloading ↓ · → · ↓ · → · ↓; ageing →↘ · ↓ (animal data only) · → · → · →↘; injury ↑ · ↓ · ↑ · ↑ · →↘ (tendinopathy only).

Mechanisms

Measured (in the human studies cited): the adult Achilles core carries 14C from childhood and adolescence; stiffness and cross-section rose with high-strain loading; stiffness fell within 1-2 weeks of limb unloading without a change in size; collagen synthesis fell by 80% after 2-3 weeks of immobilisation.
Measured (animals and cells): stretch makes tendon fibroblasts release growth factors (IGF-I, TGF-β1, CTGF) that induce collagen expression; in rats loading raised collagen and IGF-I expression similarly whether the contraction was concentric, isometric or eccentric; in 6-month-old mice overload hypertrophy came from the most superficial layers while the original core stayed relatively constant.
Authors' interpretation: tendon consists of a large stable pool formed during growth and a small fast-turnover pool for daily maintenance; it "may be speculated" that loading-induced growth adds new matrix at the periphery; high-strain tendons such as the Achilles may "not afford" constant remodelling and may turn over more slowly than the patellar tendon; the rapid loss of stiffness without loss of size may relate to cross-links formed by lysyl oxidase.
Authors' interpretation: inflammation is needed for the acute collagen response to exercise (blocking it removed the rise), distinct from the inhibiting effect of chronic inflammation.
Inference: the clock is asymmetric. Limb unloading or immobilisation costs stiffness within one or two weeks, while the gains in the training studies cited came over many weeks. Those were studies of a limb taken off weight-bearing, far more extreme than a break from running with normal walking; whether a training break alone lowers tendon stiffness, and how fast it returns, is not timed in this review.

Practical Application

Why does my Achilles lag behind my fitness when I start running? Its core barely renews after adolescence; it adapts to high strain more than moderate, and loses stiffness in 1-2 weeks of limb unloading. Ramp load slowly (practice).
High loads, any contraction type: in healthy Achilles tendons loading at about 90% of maximal contraction (about 5% strain) raised stiffness and size more than loading at 55% did, and concentric and eccentric training gave similar hypertrophy (B+, the review's summary of small human studies; the base's entry for that trial: REF-1153).
Disuse is quick to cost stiffness: 1-2 weeks of limb unloading or immobilisation lowered tendon stiffness without a change in size (human studies cited). After an injury lay-off on crutches or in a boot, expect the tendon to need re-loading before full running volume (inference; a normal training break was not studied). Not from this paper; the base's own practice for ramping load is in SYNTH-29 and SYNTH-75.
Age alone need not weaken the tendon: the lower tendon modulus of older people vanished when activity was matched (one cross-sectional study cited); keeping load in a masters athlete's week maintains the tendon (REF-599).
After a rupture, think in months: a repaired Achilles still showed raised metabolic activity at 12 months, and the authors put normalisation sometime after 6-12 months; return decisions belong to the clinician (SYNTH-73).
What it is good for as a source: the slow renewal of the tendon core, the strain dependence of adaptation, and the fast loss of stiffness with unloading. What it does not show: a weekly progression rate, a time course for stiffness gains, or injury rates; the authors note that recovery between loading sessions has not been studied.

Limitations

Narrative review. No search method or appraisal; statements are the authors' synthesis, and many draw on the Copenhagen group's own studies.
Animal and cell evidence for the cellular story. The authors themselves say the response of adult human tendon to acute loading "does not seem to mimic that of rodent tendon tissue", and that rodent loading studies use still-growing animals.
Thin human data on structure. The fibril data are "relatively scant cross-sectional" and "inconclusive"; the training evidence for stiffness comes from a few small studies; no time course of stiffness gains is given.
A detail that does not add up: the text describes tendinopathy with "cell accumulation" and "increased cell number", while the summary table marks cell number in injury as falling (↓).
"Not renewed after 17 years of age" is stated more firmly here than in the bomb-pulse paper itself, where 17 years is a model assumption based on height growth and renewal is "very limited" (REF-1356).
Funding and interests: Danish Council for Independent Research, Lundbeck Foundation and the Center for Healthy Aging (as printed; metadata add the Nordea and Novo Nordisk Foundations); competing interests none declared. No device maker, federation or data platform is named.
How this entry was checked and changed
Why this level

an authoritative narrative review from the leading group on human tendon physiology (Institute of Sports Medicine Copenhagen), read from the full text; no systematic method, and much of the cellular and molecular evidence is from animals and cell cultures, while the human loading evidence it relies on is a few small training studies and cross-sectional comparisons (on fibril structure the authors call the human data "relatively scant" and "inconclusive")

Full text (checked 2026-10-28)
Some studies in humans have found relatively high collagen synthesis rates in tendons of ∼1% per 24 h, while others have found more moderate values of around 0.2% per 24 h (full text)
The half-life of horse tendon is around 200 years in high strain superficial digital flexor tendon and ∼34 years in low strain common digital extensor tendon (full text)
Data obtained using the bomb pulse method indicates that the collagen structure of the core of the human Achilles tendon changes until about the age of 17 years and thereafter the tissue structure is stable (full text)
Subjects with a side‐to‐side difference (22%) in knee extensor strength also have a greater tendon cross‐sectional area (20%) on the stronger side (full text)
Exercising at 90% of maximum voluntary contraction (MVC; ∼5% tendon strain) yields increased stiffness and cross‐sectional area compared to working at 55% MVC (∼3% tendon strain); long duration loading (6 s cycle) more than 2 s cycle (full text)
In healthy subjects 12 weeks of resistance training comprising either concentric or eccentric knee extensions produced similar magnitudes of tendon hypertrophy (full text)
Even 2–3 weeks of immobilization of a lower limb in both young and old individuals will lead to an 80% reduction in synthesis rate of collagen (full text)
Rather short‐term (1–2 weeks) limb unloading seems to significantly reduce the stiffness of the tendon, in the absence of any change in cross‐section (full text)
Old subjects (66 years) had a lower modulus compared to young subjects (26 years), but when matched for activity level there was no difference in mechanical properties (full text)
Glucose uptake is higher in repaired than in intact Achilles tendons at 3 months (6x), 6 months (3x) and even 12 months (1.6x) after a surgical repair; not normalized until sometime after 6–12 months (full text)
The work was supported by the Danish Council for Independent Research, Lundbeck Foundation and the Center for Healthy Aging; competing interests none declared (full text)