Sir Charles S. Sherrington (1906) was one of the first to recognize the complexity
of spinal reflex circuits. Autogenic reflex pathways influence the muscles from
which they arise whereas heterogenic pathways influence the activation of
muscles apart from the muscles of origin. Both types of pathways are also
influenced by supra-spinal structures. To effectively comment on the significance
of these pathways, data on their neural interconnectivity has to be combined with
the functional consequences using biomechanical data such as the resulting
changes in joint stiffness.
Joint stiffness is made up of the algebraic sum of the stiffness of all muscles and
ligaments crossing it. The muscle stiffness constitutes passive, intrinsic and reflex
stiffness. However, in the execution of a movement, several joints are typically
displaced and it was initially proposed by Winter (1981) that dynamic balance is
maintained during standing, walking and running as the total leg stiffness never
varies. The contribution made by the reflex stiffness to the total joint stiffness has
been quantified only for the ankle joint of the lower limb where it attains values of
up to 50%. Similar data is not available for the knee joint.
This PhD project was designed to investigate the functional significance and the
neural pathways underlying the human quadriceps stretch reflex; in particular the
following aspects were studied: i) the reflex and non-reflex stiffness of the
quadriceps muscles quantified by the methods developed for the ankle joint
(Sinkjær et al., 1988), ii) the integration of the afferent information arising from
the stretch reflex at the spinal and cortical level, iii) how the central nervous
system makes functional use of this afferent information. The stretch reflex was
elicited by a whole joint rotation using either a stationary or a portable stretch
device. Subjects without a prior history of neurological diseases participated in the
various studies presented in this thesis. Such data will also be important in the
artificial control of the quadriceps during FES assisted standing in paraplegic
patients.
The results demonstrate that the human quadriceps stretch reflex can contribute
importantly to the total knee joint stiffness. Electromyographic (EMG) recordings
revealed that the response is comprised of at least three burst similar to the soleus
stretch reflex. These burst were labeled short (SLR), medium (MLR) and late
(LLR) latency reflex. While the mechanical response is similar for ankle and knee
extensors, this was not always the case for the neural responses observed. For
example, while the SLR of the soleus is strongly enhanced during co-contraction
compared to isolated plantar flexor activity, the SLR of the quadriceps is
decreased during co-contraction. Such discrepancies may be reflective of the
monosynaptic and polysynaptic heteronymous projections to quadriceps
motoneurons. The quadriceps motoneurons thus receive substantial excitatory and
inhibitory inputs from both their antagonists as well as the muscles spanning the
ankle joint (Marque et al., 2001; Marque, et al. 2005). Such connections have
been postulated to reflect a functional coupling between muscles acting as
synergists. The studies presented in this thesis provide important information on
the role of afferent feedback during an imposed stretch to a single joint and
muscle group. However the results also emphasize the need to further investigate
the role of heteronymous reflex pathways as these provide the ability to link
muscles that span different joints.