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author:("Ge, weihing")
1.  Using vertebral movement and intact paraspinal muscles to determine the distribution of intrafusal fiber innervation of muscle spindle afferents in the anesthetized cat 
Increasing our knowledge regarding intrafusal fiber distribution and physiology of paraspinal proprioceptors may provide key insights regarding proprioceptive deficits in trunk control associated with low back pain and lead to more effective clinical intervention. The use of vertebral movement as a means to reliably stretch paraspinal muscles would greatly facilitate physiological study of paraspinal muscle proprioceptors where muscle tendon isolation is either very difficult or impossible. The effects of succinylcholine (SCh) on 194 muscle spindle afferents from lumbar longissimus or multifidus muscles in response to computer-controlled, ramp and hold movements of the L6 vertebra were investigated in anesthetized cats. Paraspinal muscles were stretched by moving the L6 vertebra 1.5 to1.7mm in the dorsal-ventral direction. Initial frequency (IF), dynamic difference (DD), their changes (Δ) following SCh injection (100-400μg ·kg−1), and post-SCh dynamic difference (SChDD) were measured. Muscle spindle intrafusal fiber terminations were classified as primary or secondary fibers as well as bag1 (b1c), bag2 (b2c), b1b2c, or chain (c) fibers. Intrafusal fiber subpopulations were distinguished using logarithmic transformation of SChDD and ΔIF distributions as established by previous investigators. Increases in DD indicate strength of b1c influence while increases in IF indicate strength of b2c influence. Out of 194 afferents, 46.9% of afferents terminated on b2c fibers, 46.4% on b1b2c fibers, 1% on b1c fibers and 5.7% terminated on c fibers. Based on these intrafusal fiber subpopulation distributions, controlled vertebral movement can effectively substitute for direct tendon stretch and allow further investigation of paraspinal proprioceptors in this anatomically complex body region.
PMCID: PMC3578157  PMID: 23229776
Muscle Spindle; Neurophysiology; Paraspinal Muscles; Trunk; Lumbar Spine; Succinylcholine
2.  The decreased responsiveness of lumbar muscle spindles to a prior history of spinal muscle lengthening is graded with the magnitude of change in vertebral position 
In the lumbar spine, muscle spindle responsiveness is affected by the duration and direction of a lumbar vertebra’s positional history. The purpose of the present study was to determine the relationship between changes in the magnitude of a lumbar vertebra’s positional history and the responsiveness of lumbar muscle spindles to a subsequent vertebral position and subsequent vertebral movement. Neural activity from multifidus and longissimus muscle spindle afferents in deeply anesthetized cats was recorded while creating positional histories of the L6 vertebra. History was induced using a displacement-controlled feedback motor. It held the L6 vertebra for 4 seconds at an intermediate position (hold-intermediate at 0mm) and at 7 positions from 0.07 to 1.55mm more ventralward and dorsalward which lengthened (hold-long) and shortened (hold-short) the lumbar muscles. Following the conditioning hold positions, L6 was returned to the intermediate position. Muscle spindle discharge at this position and during a lengthening movement was compared between hold-intermediate and hold-short conditionings and between hold-intermediate and hold-short conditionings. We found that regardless of conditioning magnitude, the 7 shortening magnitudes similarly increased muscle spindle responsiveness to both vertebral position and movement. In contrast, the 7 lengthening magnitudes produced a graded decrease in responsiveness to both position and movement. The decrease to position became maximal following conditioning magnitudes of ~0.75 mm. The decrease to movement did not reach a maximum even with conditioning magnitudes of ~1.55 mm. The data suggest that the fidelity of proprioceptive information from muscle spindles in the low back is influenced by small changes in the previous length history of lumbar muscles.
PMCID: PMC3749777  PMID: 22721784
lumbar spine; muscle spindle; proprioception; thixotropy; multifidus; longissimus
Background Context
In neutral spinal postures with low loading moments the lumbar spine is not inherently stable. Small compromises in paraspinal muscle activity may affect lumbar spinal biomechanics. Proprioceptive feedback from muscle spindles is considered important for control of muscle activity. Because skeletal muscle and muscle spindles are thixotropic, their length history changes their physical properties. The present study explores a mechanism that can affect the responsiveness of paraspinal muscle spindles in the lumbar spine.
This study had two aims: to extend our previous findings demonstrating the history dependent effects of vertebral position on the responsiveness of lumbar paraspinal muscle spindles; and to determine the time course for these effects. Based upon previous studies, if a crossbridge mechanism underlies these thixotropic effects, then the relationship between the magnitude of spindle discharge and the duration of the vertebral position will be one of exponential decay or growth.
Study Design/Setting
A neurophysiological study using the lumbar spine of a feline model.
The discharge from individual muscle spindles afferents innervating lumbar paraspinal muscles in response to the duration and direction of vertebral position were obtained from teased filaments in the L6 dorsal roots of 30 Nembutal-anesthetized cats. The L6 vertebra was controlled using a displacement-controlled feedback motor and was held in each of 3 different conditioning positions for durations of 0, 0.5, 1, 1.5, and 2 seconds. Two of the conditioning positions stretched or shortened the lumbar muscles relative to an intermediate conditioning position. Conditioning positions for all cats ranged from 0.9 – 2.0 mm dorsal and ventralward relative to the intermediate position. These magnitudes were determined based upon the displacement that loaded the L6 vertebra to 50–60% of the cat’s body weight. Conditioning was thought to simulate a motion segment’s position that might be passively maintained due to fixation, external load, a prolonged posture, or structural change. Following conditioning positions that stretched (hold-long) and shortened (hold-short) the spindle, the vertebra was repositioned identically and muscle spindle discharge at rest and to movement was compared with conditioning at the intermediate position.
Lumbar vertebral positions maintained for less than 2 seconds were capable of evoking different discharge rates from lumbar paraspinal muscle spindles despite the vertebra having been returned to identical locations. Both resting spindle discharge and their responsiveness to movement were altered. Conditioning vertebral positions that stretched the spindles decreased spindle activity and positions that unloaded the spindles increased spindle activity upon returning the vertebra to identical original (intermediate) positions. The magnitude of these effects increased as conditioning duration increased to 2 seconds. These effects developed with a time course following a first order exponential reaching a maximal value after approximately 4 seconds of history. The time constant for a hold-short history was 2.6 seconds and for a hold-long history was approximately half of that at 1.1 seconds.
Thixotropic contributions to the responsiveness of muscles spindles in the low back are caused by the rapid, spontaneous formation of stable crossbridges. These sensory alterations due to vertebral history would represent a proprioceptive input not necessarily representative of the current state of intersegmental positioning. As such, they would constitute a source of inaccurate sensory feedback. Examples are presented suggesting ways in which this novel finding may affect spinal physiology.
PMCID: PMC2366132  PMID: 17938002
Background Context
Spinal manipulation (SM) is a form of manual therapy used clinically to treat patients with low back and neck pain. The most common form of this maneuver is characterized as a high velocity (duration < 150ms), low amplitude (segmental translation < 2mm, rotation < 4°, and applied force 220-889N) impulse thrust (HVLA-SM). Clinical skill in applying an HVLA-SM lies in the practitioner's ability to control the duration and magnitude of the load (i.e., the rate of loading), the direction in which the load is applied, and the contact point at which the load is applied. Control over its mechanical delivery presumably related to its clinical effects. Biomechanical changes evoked by an HVLA-SM are thought to have physiological consequences caused, at least in part, by changes in sensory signaling from paraspinal tissues.
If activation of afferent pathways does contribute to the effects of an HVLA-SM, it seems reasonable to anticipate that neural discharge might increase or decrease in a non-linear fashion as the thrust duration thrust approaches a threshold value. We hypothesized that the relationship between the duration of an impulsive thrust to a vertebra and paraspinal muscle spindle discharge would be non-linear with an inflection near the duration of an HVLA-SM delivered clinically (<150ms). In addition, we anticipated that muscle spindle discharge would be more sensitive to larger amplitude thrusts.
Study Design/Setting
A neurophysiological study of spinal manipulation using the lumbar spine of a feline model.
Impulse thrusts (duration: 12.5, 25, 50, 100, 200, and 400 ms; amplitude 1 or 2mm posterior to anterior) were applied to the spinous process of the L6 vertebra of deeply anesthetized cats while recording single unit activity from dorsal root filaments of muscle spindle afferents innervating the lumbar paraspinal muscles. A feedback motor was used in displacement control mode to deliver the impulse thrusts. The motor's drive arm was securely attached to the L6 spinous process via a forceps.
As thrust duration became shorter the discharge of the lumbar paraspinal muscle spindles increased in a curvilinear fashion. A concave up inflection occurred near the 100ms duration eliciting both a higher frequency discharge compared to the longer durations and a substantially faster rate of change as thrust duration was shortened. This pattern was evident in paraspinal afferents with receptive fields both close and far from the midline. Paradoxically, spindle afferents were almost twice as sensitive to the 1mm compared to the 2mm amplitude thrust (6.2 vs 3.3 spikes/s/mm/s). This latter finding may be related to the small vs large signal range properties of muscle spindles.
. The results indicate that the duration and amplitude of a spinal manipulation elicits a pattern of discharge from paraspinal muscle spindles different from slower mechanical inputs. Clinically, these parameters may be important determinants of an HVLA-SM's therapeutic benefit.
PMCID: PMC2075482  PMID: 17905321
lumbar spine; spinal manipulation; chiropractic; osteopathy; paraspinal muscles; muscle spindle

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