Operant Conditioning of H-Reflex Can Correct a Locomotor Abnormality after Spinal Cord Injury in Rats

Yi Chen, Xiang Yang Chen, Lyn B. Jakeman, Lu Chen, Bradford T. Stokes, Jonathan R. Wolpaw · Journal of Neuroscience · 2006 · neuroscience · 10.1523/jneurosci.2198-06.2006 · record chen2006-operant-conditioning-of-h-reflex-c
tag layer · index · concepts · authors · עברית
Where this record's text came from. faceted 2026-08 by reading the locally held text (html); every evidence string verified as an exact substring of that text before merging. method tagged 'electrophysiology' because the measurements are nerve-cuff-evoked H-reflexes and chronic EMG; the design is equally an animal behavioural/operant-conditioning experiment with a lesion manipulation, which the single-value facet cannot express. n is small (8 HRup, 5 control rats), and the paper reports this openly but does not frame it as a limitation, so it is not in limitations. Source is a PMC HTML dump: inline citations carry extra spaces inside the parentheses (e.g. '( Chen et al., 2005 )') and quotes preserve that spacing verbatim. Boilerplate .gov/PMC navigation text at the top of the file was ignored. 2026-08-13 and 2026-08-15 schema-0.3 passes: every quote verified as an exact substring of the staged text and anchored at merge time; concept terms verified present; every edge endpoint an existing node.
How to read this page. The article itself is not reproduced here — this paper is not cleared for redistribution, so what follows is the tag layer alone. The evidence sentences are withheld with the article. The table at the foot lists every tag the record carries; each row names the facet it belongs to, and the whose words column says whether the tag records the paper's own wording or the tagger's sentence about the paper. Every tag links to each other paper in the corpus carrying it. The machine-readable edition of the same thing is the TEI file beside this one, where the tags live in standOff, and the article is at its publisher via the link below. When we have changed our mind. Some entries in the table begin with a word in capitals and the reason for it. EVIDENCE CORRECTED — the finding still stands, but we had quoted the wrong sentence from the paper, and the quotation has been replaced with the right one. RETRACTED — the entry itself was wrong; it stays on the page, marked, with the reason, rather than being deleted. WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below. Each mark carries its own reason, in brackets, because the reasons differ from one entry to the next. An entry we have WITHDRAWN is removed from this page and kept in the record, with its reason. A line reading Audit and a date is a later re-reading of the entry against its whole sentence: DOUBTFUL means it was kept, and the reason says what is loose about it. Quotation marks inside the tagger's sentence. Where a tag is the tagger's own sentence, a phrase in quotation marks inside it is borrowed from the paper — the tagger's wording around it is not. The paper's own sentence is the evidence quote for that row; on this page it is withheld with the article, and the article is at its publisher. Nothing inside quotation marks anywhere on this page is ours: if it is quoted, it is the paper's. Whose words a concept carries. A concept entry records a term the paper itself uses, so the term, the sense given for it, and its evidence are the paper's own words, quoted where the paper fixes them. This is the opposite of a sense entry in the tag table, which is the tagger's sentence about how the paper uses a term. Where a concept entry could not be grounded in the paper's words, the entry says so on its own row.
Other fields define these differently the same word, defined another way in another discipline
coefficient of variation 4 fields · standard deviation 3 fields

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 12 concept tags on this paper, none is grounded in a quoted sentence. All carry the tagger's gloss of what the term means in this paper, which is not something the article can license.
H-reflex operant conditioning spinal cord plasticity lateral column corticospinal tract locomotor asymmetry step-cycle symmetry soleus BBB score M-response background EMG stance phase Up-conditioning strengthens the motoneuron response to primary afferent input, recruiting more motoneurons into the locomotor burst (and making each fire more), which enlarges and broadens the soleus burst and thereby lengthens right stance until symmetry is restored The enlarged locomotor burst produced by up-conditioning is the proposed origin of the restored right/left symmetry In normal animals compensatory adjustments by other muscles cancel the conditioning-induced change; in lesioned animals those compensatory adjustments are impaired or absent, so the conditioning effect is allowed to alter locomotion spinal cord injury locomotion treadmill step cycle asymmetry EMG motoneuron stance up-conditioning down-conditioning rehabilitation nerve cuff conditioning H-reflex successful conditioning burst onset burst offset H-reflex up-conditioning treadmill speed M-response size H-reflex size burst amplitude burst width ANOVA one-way repeated-measures ANOVA post hoc multiple comparison (Tukey's test) pairwise multiple comparison (Tukey's test) t-test t test correlation standard deviation mean ± SD coefficient of variation one-way repeated-measures ANOVA with Tukey's test coefficient of variation across step cycles electrophysiology

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second readingalternativesan independent re-reading of a facet, recorded whether or not it agrees
relation to other workclaim_relationshow this paper stands to another: agrees, contradicts, extends, qualifies
claim typeclaim_typethe shape of the claim: existence, mechanism, method, critique, prediction, null
concept edgeconcept_edgesan asserted relation between two concepts, with the sentence asserting it
content tagcontent_tagsa finding, definition, quantity, statistic or variable named in the paper
contributioncontributionwhat the paper claims to add
controlcontrolsan experiment run to eliminate a rival explanation
limitationlimitationswhat the paper says its evidence cannot support
mechanismmechanismshow the reported effect is said to come about
methodmethodhow the paper produced its evidence
object of studyobject_of_studywhat the paper is about — the thing it investigates
phenomenonphenomenonthe effect or regularity under study
research questionresearch_questionthe question the paper sets itself
sensesensesa distinct use of a term, quoted where the paper fixes it
transfertransfersa method or concept carried in from another field

The paper

The full text is not reproduced here. This article is not cleared for redistribution, so the page carries the tag layer only. Its evidence sentences are withheld; the grounding status remains with each tag, and the article itself is at https://doi.org/10.1523/jneurosci.2198-06.2006. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 3 · claim_relations 6 · claim_type 1 · concept_edges 5 · content_tags 52 · contribution 1 · controls 1 · limitations 12 · mechanisms 8 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 4 · transfers 2

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 5 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studywhether operant conditioning of a spinal reflex can change locomotion after spinal cord injury
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object_of_studythe paper's words
research_questionwhether descending input shaped by reflex conditioning can improve motor function in the injured spinal cord
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research_questionthe paper's words
contributionshows a targeted reflex conditioning protocol removes a lesion-induced locomotor asymmetry, establishing the principle that descending-input conditioning can change a complex motor behaviour
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contributionthe paper's words
claim_typeexistence
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claim_typethe paper's words
phenomenonH-reflex operant conditioning after spinal cord injury
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phenomenonthe paper's words
methodelectrophysiology
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methodthe paper's words
mechanismsmechanisms
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mechanisms[0]the tagger's words
mechanismsmechanisms
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mechanisms[1]the tagger's words
mechanismsmechanisms
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mechanisms[2]the tagger's words
mechanismsup-conditioning raises the locomotor burst, probably by recruiting more motoneurons to primary afferent input, and the burst also broadens
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mechanisms[3]0.95the tagger's words
mechanismsthe lesion spares the corticospinal tract, which is the only descending path known to be essential for H-reflex conditioning - so the protocol can still work
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mechanisms[4]1.0the tagger's words
mechanismsin intact animals other muscles compensate and symmetry is preserved; after injury that compensation is impaired, so the same conditioning changes locomotion
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mechanisms[5]1.0the tagger's words
mechanismsconditioning is delivered continuously in the home cage, 3000-9000 stimulus trials per day over 50 days
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mechanisms[6]0.8the tagger's words
mechanismsH-reflex size is expressed in units of background EMG rather than peak-to-peak, because the rat H-reflex form is too variable for a peak measure
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mechanisms[7]1.0the tagger's words
limitationsThe study only establishes the principle; the functional and clinical significance of the effect is not yet assessed.
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what this limits: what the study establishes, stated by the authors as principle only: THE PRESENT RESULTS MERELY VALIDATE THE PRINCIPLE THAT SPINAL REFLEX CONDITIONING CAN AFFECT A COMPLEX MOTOR FUNCTION AFTER SPINAL CORD INJURY
limitations[0]the tagger's words
limitationsWhether treadmill results in quadrupedal rats transfer to over-ground walking in bipedal humans is unknown.
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what this limits: transfer to human walking: THE EXTENT TO WHICH THE EFFECTS OF CONDITIONING ON TREADMILL LOCOMOTION IN THE QUADRIPEDAL RAT PREDICT ITS EFFECTS ON OVER-GROUND LOCOMOTION IN THE BIPEDAL HUMAN REMAINS TO BE DETERMINED
limitations[1]the tagger's words
limitationsHow long the conditioning effects last is not established.
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what this limits: any claim about how long the effect lasts: THE LONG-TERM PERSISTENCE OF THE EFFECTS OF H-REFLEX CONDITIONING IS NOT YET WELL DEFINED
limitations[2]the tagger's words
limitationsLarge between-rat variability in absolute EMG measures limits the power to detect group differences.
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what this limits: between-group comparisons on absolute measures: THIS WIDE INTER-RAT VARIABILITY MEANS THAT VERY LARGE GROUPS OF LC AND NORMAL RATS WOULD BE NEEDED FOR INTERGROUP DIFFERENCES TO BECOME SIGNIFICANT
limitations[3]the tagger's words
limitationsthe measured asymmetry is an EMG proxy, and whether it corresponds to an actual limp is stated to be unknown
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what this limits: the step from the EMG measure to the behaviour: IT IS NOT AS YET CLEAR WHETHER THE SOLEUS EMG ASYMMETRY CREATED BY THE LC LESION IS ASSOCIATED WITH AN ACTUAL LIMP IN WALKING
limitations[4]1.0the tagger's words
limitationsthe results are said to merely validate a principle, with clinical significance unassessed
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what this limits: the same scope as limitations[0] - the principle, not the clinical application: MUCH REMAINS TO BE LEARNED BEFORE THE POTENTIAL of the method is known
limitations[5]1.0the tagger's words
limitationsonly soleus bursts were measured; stance and swing need direct measurement and other muscles are untested
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what this limits: the soleus-burst proxy for stance and swing: THE DATA PRESENTED HERE RELY ON THE RIGHT AND LEFT SOLEUS LOCOMOTOR BURSTS TO ASSESS SPECIFIC ASPECTS OF LOCOMOTION, and these measures only APPEAR TO PARALLEL THE RIGHT AND LEFT STANCE PHASES OF LOCOMOTION
limitations[6]1.0the tagger's words
limitationswhether quadrupedal treadmill results predict bipedal over-ground human locomotion is undetermined
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duplicate of: limitations[1] · what this limits: the same scope as limitations[1] - prediction from quadrupedal treadmill work to BIPEDAL HUMAN over-ground locomotion
limitations[7]1.0the tagger's words
limitationsthe long-term persistence of the effect is not well defined, and up-conditioning decays faster than down-conditioning in monkeys
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what this limits: the same scope as limitations[2] - persistence, where the human comparison is that THE EFFECTS OF DOWN-CONDITIONING ARE STILL EVIDENT 4 MONTHS LATER
limitations[8]1.0the tagger's words
limitationsinter-rat variability in absolute EMG measures is so wide that very large groups would be needed for group differences to reach significance
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what this limits: absolute measures specifically: ABSOLUTE MEASURES SUCH AS SOLEUS BURST AMPLITUDE DIFFER WIDELY ACROSS NORMAL RATS AND EVEN BETWEEN THE RIGHT AND LEFT LEGS OF INDIVIDUAL RATS
limitations[9]1.0the tagger's words
limitationsthe locomotor analysis uses only the seven rats in which conditioning succeeded
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what this limits: the up-conditioning locomotion analysis, which used only the animals in which conditioning worked: THE SEVEN SUCCESSFUL RATS CONSTITUTED THE EXPERIMENTAL GROUP OF HRup RATS FOR EVALUATION OF THE EFFECTS OF UP-CONDITIONING ON TREADMILL LOCOMOTION
limitations[10]1.0the tagger's words
limitations13 female rats only
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what this limits: any sex-general reading: SUBJECTS WERE 13 FEMALE SPRAGUE DAWLEY RATS
limitations[11]0.85the tagger's words
transfersreward-contingent shaping, applied to a spinal reflex rather than to a voluntary action
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transfers[0]1.0the tagger's words
transfersevidence that far fewer trials suffice in people, used to argue the rat protocol's intensity is not a barrier
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transfers[1]1.0the tagger's words
controlsfive rats that were never exposed to up-conditioning, run through the same two sessions as the conditioned animals
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eliminates: the passage of time and repeated testing as the explanation of the change - if the unconditioned rats changed too, the effect would not be the conditioning · outcome: held
controls[0]0.95the tagger's words
senseslocomotor abnormality here means an EMG burst-onset asymmetry, not an observed gait defect - the paper says whether there is a limp at all is unknown
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senses[0]1.0the tagger's words
sensessuccessful conditioning is stipulated as a change of at least 20% in the intended direction, a criterion carried over from earlier papers
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senses[1]1.0the tagger's words
sensesthe abnormality is PHYSIOLOGICALLY defined and the lesion ANATOMICALLY defined - the paper states the two bases separately
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senses[2]1.0the tagger's words
sensessymmetry is an index constructed so that 0.5 means symmetric, built from burst-onset times rather than from limb kinematics
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senses[3]1.0the tagger's words
alternativesclaim_type: mechanism
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a second reading DIFFERING from the recorded value. Recorded: existence — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.claim_typethe tagger's words
alternativesmethod: right lateral column transection, then 50 days of H-reflex up-conditioning by nerve-cuff stimulation and food reward in the freely moving rat, with treadmill locomotion measured before and after
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a second reading DIFFERING from the recorded value. Recorded: electrophysiology — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.methodthe tagger's words
alternativesphenomenon: activity-dependent spinal cord plasticity
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a second reading DIFFERING from the recorded value. Recorded: H-reflex operant conditioning after spinal cord injury — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationsextends → Chen et al., 2005 (H-reflex conditioning in normal rats)
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claim_relations[0]the tagger's words
claim_relationsbuilds-on → Chen et al., 2001a and Chen et al., 2005
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claim_relations[1]the tagger's words
claim_relationsextends → work showing that patterned sensory input from locomotor training induces spinal plasticity, by adding a descending-input route
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claim_relations[2]1.0the tagger's words
claim_relationssupports → the feasibility of reflex conditioning in people with partial spinal cord injuries who retain a functioning CST
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claim_relations[3]0.95the tagger's words
claim_relationsqualifies → its own translational claim, by stating the rat-to-human locomotion question is undetermined
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status: the paper's own words: remains to be determined
claim_relations[4]1.0the tagger's words
claim_relationsdistinguishes-from → the same conditioning in normal rats, where it does not change locomotor symmetry at all
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differs in: THE AXIS IS WHETHER THE ANIMAL IS INJURED. 'Quotation withheld' The same conditioning does nothing in an intact animal and changes locomotion in an injured one - so the effect is a property of the DAMAGED system, and a therapy claim depends entirely on that.
claim_relations[5]1.0the tagger's words
concept_edgesH-reflex —causes→ locomotor asymmetry
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strength of the relation: 0.85 · note: SCOPE: this holds in lesioned rats and explicitly does NOT hold in normal rats, where the same conditioning leaves symmetry unchanged
concept_edges[1]1.0the tagger's words
concept_edgeslateral column —distinct-from→ corticospinal tract
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strength of the relation: 1 · note: the whole design rests on this separation - the lesion removes one and spares the other Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the procedure transected the LC and left the CST intact; distinct-from is loose.
concept_edges[2]1.0the tagger's words
concept_edgeslocomotor asymmetry —operationalised-as→ step-cycle symmetry
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span defines step-cycle symmetry; 'locomotor asymmetry' is not named.
concept_edges[3]1.0the tagger's words
concept_edgesEVIDENCE CORRECTED (RE-GROUNDED) · H-reflex —operationalised-as→ background EMG
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EVIDENCE CORRECTED: RE-GROUNDED. The old span explained why the measure is used. This sentence IS THE OPERATIONALISATION, as a formula: '…and was expressed in units of average background EMG…'. The second endpoint appears twice - as the subtrahend and as the unit. — strength of the relation: 0.9 · note: the measurement unit was chosen because the animal's reflex form is too variable for the obvious measure Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but endpoint by anaphora: 'in this manner' = measured relative to average background EMG amplitude (previous sentence).
concept_edges[4]1.0the tagger's words
concept_edgesH-reflex —operationalised-as→ background EMG
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strength of the relation: 0.9 · note: RE-GROUNDS another row of this record: the old span named ONE endpoint. Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but H-reflex size is expressed IN UNITS of background EMG; operationalised-as is loose.
concept_edges[5]1.0the tagger's words
content_tagsANOVA
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also in: ANOVA
content_tags.statistics[0]the tagger's words
content_tagspost hoc multiple comparison (Tukey's test)
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content_tags.statistics[1]the tagger's words
content_tagst-test
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also in: t-test
content_tags.statistics[2]the tagger's words
content_tagscorrelation
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also in: correlation
content_tags.statistics[3]the tagger's words
content_tagsstandard deviation
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content_tags.statistics[4]the tagger's words
content_tagscoefficient of variation
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content_tags.statistics[5]the tagger's words
content_tagsone-way repeated-measures ANOVA with Tukey's test
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content_tags.statistics[6]0.8the tagger's words
content_tagscoefficient of variation across step cycles
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content_tags.statistics[7]0.85the tagger's words
content_tagsUp-conditioning the soleus H-reflex enlarged the right soleus burst and corrected the locomotor asymmetry.
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content_tags.findings[0]the tagger's words
content_tagsRight lateral-column transection shortened the right stance phase: right-to-left burst-onset time was only 77% of left-to-right.
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content_tags.findings[1]the tagger's words
content_tagsThe conditioning protocol roughly doubled the size of the conditioning H-reflex while leaving background EMG and the direct muscle (M) response unchanged, showing the change was reflex-specific rather than a general increase in muscle activation.
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content_tags.findings[2]the tagger's words
content_tagsIn up-conditioned rats the locomotor H-reflexes and right soleus burst amplitude and width grew significantly between the first and second sessions.
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content_tags.findings[3]the tagger's words
content_tagsStep-cycle duration did not change in either the up-conditioned or the control group.
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content_tags.findings[4]the tagger's words
content_tagsIn normal rats neither up- nor down-conditioning affects right/left step symmetry.
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content_tags.findings[5]the tagger's words
content_tagsUp-conditioning succeeded in 7 of 8 lesioned rats (87.5%).
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content_tags.findings[6]the tagger's words
content_tagslateral column transection left a persistent burst-onset asymmetry: right-to-left time was 0.43 of the cycle against 0.50 in normal rats
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content_tags.findings[7]1.0the tagger's words
content_tagsthe right stance phase was shortened to 77% of the opposite interval
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content_tags.findings[8]1.0the tagger's words
content_tagsup-conditioning succeeded in 7 of 8 rats
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content_tags.findings[9]1.0the tagger's words
content_tagslocomotor H-reflexes and right soleus burst amplitude and width all increased in the conditioned group
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content_tags.findings[10]1.0the tagger's words
content_tagsthe asymmetry disappeared in the up-conditioned group
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content_tags.findings[11]1.0the tagger's words
content_tagsa stance H-reflex increase came with a burst amplitude increase 0.32 times as large
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content_tags.findings[12]1.0the tagger's words
content_tagsthe same conditioning does not change locomotor symmetry in normal rats
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content_tags.findings[13]1.0the tagger's words
content_tags72% of the right lateral column was destroyed, with other structures mostly intact
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content_tags.findings[14]1.0the tagger's words
content_tagsconditioning H-reflex
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content_tags.definitions[0]the tagger's words
content_tagsstep-cycle symmetry
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content_tags.definitions[1]the tagger's words
content_tagssuccessful conditioning
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content_tags.definitions[2]the tagger's words
content_tagsstep-cycle symmetry
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content_tags.definitions[5]1.0the tagger's words
content_tagssuccessful conditioning
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content_tags.definitions[6]1.0the tagger's words
content_tagsburst onset
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also in: burst onset
content_tags.definitions[7]the tagger's words
content_tagsburst offset
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also in: burst offset
content_tags.definitions[8]the tagger's words
content_tagsH-reflex up-conditioning
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content_tags.variables[0]the tagger's words
content_tagsstep-cycle symmetry
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content_tags.variables[1]the tagger's words
content_tagstreadmill speed
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content_tags.variables[3]the tagger's words
content_tagsM-response size
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content_tags.variables[4]the tagger's words
content_tagsH-reflex size
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also in: H-reflex size
content_tags.variables[5]1.0the tagger's words
content_tagsburst amplitude
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content_tags.variables[8]1.0the tagger's words
content_tagsburst width
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also in: burst width
content_tags.variables[9]1.0the tagger's words
content_tags212 ± 49% — final size of the conditioning H-reflex in successfully up-conditioned rats, as a percentage of its pre-conditioning control size
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content_tags.quantities[0]the tagger's words
content_tags87.5% — proportion of rats (7 of 8) in which up-conditioning met the success criterion
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content_tags.quantities[1]the tagger's words
content_tags0.43 ± 0.02 — right-to-left burst-onset interval as a fraction of the whole step cycle in lesioned rats, versus the 0.50 of symmetrical locomotion
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content_tags.quantities[2]the tagger's words
content_tags280 ± 38% — increase in the stance-phase locomotor H-reflex after up-conditioning, as a percentage of its first-session value
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content_tags.quantities[3]the tagger's words
content_tags158 ± 22% — increase in right soleus locomotor burst amplitude after up-conditioning, as a percentage of its first-session value
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content_tags.quantities[4]the tagger's words
content_tags72 ± 18% — proportion of the right lateral column destroyed by the transection, verified histologically
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content_tags.quantities[5]the tagger's words
content_tagsp = 0.008 — significance of the disappearance of the right/left step-cycle asymmetry in the up-conditioned group
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content_tags.quantities[6]the tagger's words
content_tags13 — number of rats used in the study
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content_tags.quantities[7]the tagger's words
content_tags13 female Sprague Dawley rats — the sample
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content_tags.quantities[8]1.0the tagger's words
content_tags10% of peak — the stipulated threshold defining burst onset and offset
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content_tags.quantities[9]1.0the tagger's words
content_tagsat least 20% change — the stipulated criterion for successful conditioning
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content_tags.quantities[10]1.0the tagger's words
content_tags0.43 versus 0.50 of the step cycle, p = 0.009 — the lesion-induced asymmetry against normal
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content_tags.quantities[11]1.0the tagger's words
content_tags212% of control, 7 of 8 rats, 87.5% success — the conditioning effect and its success rate
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content_tags.quantities[12]1.0the tagger's words
content_tags0.32 times as large, r = 0.60, p = 0.04 — burst amplitude change per unit H-reflex change
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content_tags.quantities[13]1.0the tagger's words
content_tags72 plus or minus 18% of the right lateral column destroyed — verified lesion extent
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content_tags.quantities[14]1.0the tagger's words

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