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Evidence that Synchronized Motor Unit Firings Are
Epiphenomena of the Onion Skin Property Not Common Inputs
Joshua C. Klinea,b and Carlo J. De Lucaa,b
a) Delsys Inc., Natick, MA, USA; b) Department of Biomedical Engineering, Boston University, Boston, MA, USA
Synchronous motor unit (MU) firing instances have been attributed to anatomical
inputs shared by motoneurons. Yet, there is a lack of empirical evidence confirming the
notion that common inputs are responsible fro synchronization during voluntary
conditions. Therefore, we designed two experiments to:
1) test the veracity of the common input notion; and
2) investigate alternative empirically-based explanations for synchronization.
References
De Luca et al, J Neurophysiol (1993)
De Luca and Contessa, J Neurophysiol (2012)
De Luca and Kline, J Neurophysiol (2014)
Motivation
Experiment 1: Testing the Common Input Notion
Subjects were required to track a two-force level paradigm that was maintained at a
relatively low force level and then increased to a relatively higher force level. We used
the dEMG System for decomposing sEMG signals into MU action potential trains
(MUAPTs) and compared the degree of synchronization measured by the SigMax
method (De Luca and Kline, 2014) between MUAPTs as their firing rates progressed
throughout the increasing force contraction.
This work was supported in part by two
grants from NIH 1R43NS077526, and a grant
from the Neuromuscular Research
Foundation.
Acknowledgements
Experiment 1: Testing The Common Input Notion
The common input notion provides no empirically supported explanation to account
for the decrease in synchronization with increasing contraction force.
Thus, the practice of inferring anatomical connections shared by motoneurons based
on observations of synchronization remains empirically unsupported.
So viewed, occasionally synchronous firing instances are likely an epiphenomenon,
subject to countless unknown neural interactions. As such, synchronization may not be
the product of a specific design and may not serve a specific physiological purpose.
Our explanation for synchronization has the advantage of being supported by
empirical evidence; whereas, the common input does not.
Experiment
Experiment 2: Investigating Empirical Relationships with Synchronization
We studied associations between synchronization and MU firing rates in different
muscles and for different voluntary contraction force levels by measuring the
synchronization index (SI) between pairs of MUAPTs active during voluntary
contractions ranging from 5% MVC to 50% MVC in the first dorsal interosseous (FDI)
and vastus lateralis (VL) muscles.
Measure Synchronization
Measure Synchronization
Lower Force Level
Higher Force Level
Compare
……
MU A
MU B
MU Mean Firing Rate
MU
Mea
n F
iring
Rat
e
Low High
Low
Hig
h
Synchronization Measurements
2
31
High Sync.
Low Sync.
No Sync.
1) SI = 30.2
2) SI = 17.5
3) SI = 0
MU Mean Firing Rates
2
31
..
.
5 10 15 200
10
20
30
40
Time (s)
Mea
n F
iring
Rat
e (p
ps)
00
5
10
15
20
25
For
ce (
% M
VC
)
-25 0 250
1
-10 0 10
0
1
Latency(ms) Latency(ms)
Pro
babi
lity
Pro
babi
lity
-10 100
We measured synchronization between the same pairs of MUAPTs as their firing rates
progressed from a contraction at a relatively low force level to a higher one. As the
contraction force increased, synchronization decreased.
FDI VL
∆ S
ync.
Inde
x (%
)
-15
0
15
10
5
-5
-10
Alt. MUAPT MFR (pps)
10 20
Alt. MUAPT MFR (pps)
10 20
10
20
Ref
. MU
AP
T M
FR
(pp
s)
∆ S
ync.
Inde
x (%
)
-15
0
15
10
5
-5
-10
Avg. MUAPT MFR (pps)
10 20
Avg. MUAPT MFR (pps)
10 20
G) ∆SI H) ∆SI
∆ S
ync.
Inde
x (%
)
-15
0
15
10
5
-5
-10
E) ∆SI < 0 (p<0.0001) F) ∆SI < 0 (p<0.0001)
Low Force
HighForce
∆SI
∆SI
Our data set of 17,546 paired MUAPTs revealed that the degree of synchronization
varies as a function of two characteristics of the firing rate: the slope and similarity of
the firing rates. Both are measures of the excitation of the motoneurons (De Luca and
Contessa, 2012). As the force generated by the muscle increases the firing rate slope
decreases and the synchronization correspondingly decreases. Different muscles have
motor units with different firing rate characteristics, and display different amounts of
synchronization.
1 1
3
2
MoreSync.
Similar firing ratesSteep slopes
ModerateSync.
LessSync.
No Sync.
Different firing rates
Similar firing ratesModerate slopes
Lower Voluntary Excitation
Higher Voluntary Excitation
Similar firing ratesShallow slopes
Firi
ng R
ate
Force
Firi
ng R
ate
Force
Firi
ng R
ate
Force
0 10 20 30 40 50 60 70
0
5
10
15
20
25
30
Time (s)
Mea
n F
iring
Rat
e (p
ps)
0
5
10
15
20
For
ce (
% M
VC
)
MU 2
MU 3
MU 14
MU 17
Experiment 2: New Explanation for Synchronization
A) SI at Lower Force B) SI at Lower Force
D) SI at Higher ForceC) SI at Higher Force
Syn
c. In
dex
(%)
0
10
20
30
Ref
. MU
AP
T M
FR
(pp
s)
10
20
Ref
. MU
AP
T M
FR
(pp
s)
10
20
Syn
c. In
dex
(%)
0
10
20
30
Avg. MUAPT MFR (pps)
10 20
Avg. MUAPT MFR (pps)
10 20
31 1
2
Synchronization Decreases with Increasing Force
“Common Input” Provides no Explanation
Synchronization is Related to the Characteristics of the
Motor Unit Firing Rates
SENSORS FOR MOVEMENT SCIENCES