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In addition, the sign of the force difference between proximal and distal EDL forces changed. Similar effects of muscle relative position were reported for slightly different experimental conditions [ 17 ]. Length-force characteristics of EDL muscle obtained by movements of the distal tendon were significantly different from its length-force characteristics if EDL muscle was lengthened by moving its proximal tendon.
In conclusion, the position of a muscle relative to surrounding tissues codetermines isometric muscle force. Position effects can be explained by changes in the configuration of the tissues representing the epimuscular myofascial pathways Figure 2. In general, the muscle end that is positioned farthest in a particular direction e. The above in situ studies have shown the potential of force transmission between skeletal muscles via inter- and extramuscular connective tissues. The functional relevance of this phenomenon is dependent on the magnitude of the effects found in physiological muscle conditions.
However, this mode of force transmission may be small in normal muscles during physiological conditions, because 1 the above studies all used tetanic stimulation. This rarely occurs during voluntary movement, so observations may be relevant only to lab conditions; 2 the muscle-tendon complex length of a single muscle was changed while the length of its synergists was kept constant, compared to simultaneous length changes in synergistic muscles during normal movements; 3 when individual muscles are stimulated alone and together, the force sums linearly which is surprising if the epimuscular pathway is used; and 4 a recent experiment studying force transmission between cat soleus SO and its synergistic muscles in an intact animal showed little epimuscular force transmission.
Each of these points will be discussed below. In the studies described up to this point, the effects of epimuscular pathways on muscular force transmission were tested predominantly during simultaneous maximal activation of both synergistic and antagonistic muscles. Coactivation of synergistic and antagonistic muscles has been observed in the awake, freely moving animal e. This suggests that also at firing frequencies encountered in vivo muscle forces can be transmitted via epimuscular myofascial pathways.
Another experimental condition of the studies described in Section 2. Due to differences in moment arms between synergists [ 2122 ], the change in length of one muscle can be different from that of its neighbor, but the relative movements imposed during lengthening a single muscle were beyond the physiological range.
It should be noted that different myofascial pathways can be arranged in such a way that they exert forces on a muscle in opposite direction see [ 9Figure ] and that the proximo-distal force difference is the net result of all myofascial loads [ 7 ]. A small difference can thus be explained by limited epimuscular myofascial force transmission or opposing myofascial loads of similar magnitude. If force transmission through epimuscular pathways is substantial, then nonlinear summation of force is expected when different muscles are activated alone and together.
Nonlinear force summation is defined as the difference between the force exerted when two muscle parts are excited simultaneously and the sum of the forces exerted when each muscle part is excited individually [ 24 ].
The muscles were activated by stimulation of the nerve branches to each of the muscle groups. LG and SO muscles were stimulated together because of the difficult surgery required to separate their nerves see [ 27 ]. The cat hindlimb was left intact and the foot attached to a 6-degree-of-freedom dof load cell to measure force and torque. The femur was fixed to a rigid frame. When both muscles were stimulated together, the resulting forces and torques all 6 dof were less than the sum of the individual forces.
There was no evidence that the direction of the forces changed during simultaneous activation of the muscles compared to activation of the muscles independently.
These results suggest that when both muscle groups were activated together there was increased shortening of the muscle fibers, and hence less force due to a higher velocity of shortening during force onset.
Thus, there is some interaction, either between the muscle bellies or between their tendons. However, the interaction was small, and during steady state, it was almost immeasurable.
Similar experiments were performed on the vastus medialis and rectus femoris in cat. Both muscles are knee extensors. They share a border and a tendon and thus may be expected to show nonlinear summation. Nonlinear summation error was small in all 6 degrees of freedom. The average peak error was 8. Note that these experiments do not preclude epimuscular force transmission, but rather suggest that in normal muscle it has little functional effect on the overall force delivered to the skeleton.
To tackle some of the concerns of previous studies, a new experimental approach was developed to measure directly the mechanical interactions between muscles in conditions that simulate those present during normal movements [ 28 ]. The latter was assured by testing the muscles in a nearly intact limb of the cat.
The tendons were not cut, but left attached to their insertion sites. Length changes were obtained by movements of the joints and, thus, only physiological relative movements could be imposed. The mechanical interactions between the one-joint SO and its two-joint synergistic muscles were studied.
Note that knee movements will only alter the length and relative position of the two-joint muscles, but not of SO. This involves the greatest relative displacements between these muscles in vivo. We hypothesized that force transmission from SO muscle fibers will be affected by length changes of its synergists through configuration changes of connective tissues between these muscles.
A schematic presentation of the cat hindlimb in the experimental setup used to investigate inter-synergist interactions [ 28 ]. Changing the knee angle from to lengthened LG and PL profoundly 4. In contrast to our expectations, active ankle moment generated by SO and the rate of muscle relaxation were not significantly affected by changes in knee angle.
These results demonstrate that the presence of relative muscle movements does not necessarily mean force transmission between muscles. To further test the apparent independency of SO, an additional set of experiments was performed. With minimal disruption of the connective tissues at the muscle belly level, the distal tendon of SO was dissected free from the other tendons in the Achilles tendon complex, cut, and connected to a force transducer.
BioMed Research International
As this eliminated force transmission to its insertion on the calcaneus, any ankle joint moment following SO excitation was attributed to force transmission via epimuscular myofascial pathways to the Achilles tendon. If the tendon of SO was placed at its original position, corresponding to the above reported ankle joint angle, the moment exerted at the ankle was close to zero while force exerted at the distal tendon of SO was near its optimal value.
A substantial ankle moment was found only if SO was excited at positions distant from physiological. These results confirm that for in vivo muscle lengths and relative positions force generated in SO muscle fibers is transmitted to its distal tendon. Others have found that human SO fascicle length was not affected by changes in knee angle, as measured in both passive and maximally active conditions of the ankle plantar flexors [ 31 ].
In contrast, recent imaging studies in humans suggest that mechanical connections between gastrocnemius and SO muscles are effective also within the in vivo length range. Isolated excitation of MG at a fixed angle of the ankle and knee joints elicited a decrease in fascicle length, not only in the excited muscle, but also in SO [ 3233 ].
However, MG activation did not cause displacements in flexor hallucis longus muscle [ 33 ], suggesting that not all muscles are equally connected.
In addition, effects of knee movements on SO muscle have been reported [ 3334 ]. Note that the mechanical effects e. How can the different results between the rat and cat studies be explained? We have hypothesized that the intermuscular linkages between SO and adjacent muscles within the intact cat may be slack or operate on the toe region of their lumped stress-strain curve Figure 4see also [ 28 ]. The steep portion of this curve and, hence, epimuscular force transmission will then be attained only with supraphysiological displacements.
The stiffness of the intermuscular linkages may also be a local property, being more compliant in the proximal region of the cat ankle extensors. Note that Maas and Sandercock [ 28 ] lengthened the two-joint muscles by knee movements; thus, there was more movement proximally. Preliminary data suggest indeed that cat SO muscle is more rigidly connected to LG distally than proximally Sandercock and Maas, unpublished observations. Also in line with this hypothesis are the results of an earlier study in the rat in which lengthening a muscle distally resulted in substantial force changes exerted at the distal tendon of a neighboring muscle, while effects of proximal lengthening were not significant [ 17 ].
Drawings to illustrate length changes of connective tissue linkages between passive grey and active red synergistic muscles. Changing the length of one muscle results in reorientation as well as unfolding of those linkages. Unfolding is also seen with coactivation. Such straightening of macroscopic crimp in collagen fibrils is correlated to the toe region of the stress-strain curve [ 35 ]. Modified from Maas and Sandercock [ 28 ].
A major difference between the experiments on epimuscular myofascial force transmission in the rat see above and the cat [ 28 ] is the number of muscles that is activated simultaneously.
In the rat studies all synergists and some antagonists were activated versus a single muscle in the cat. Coactivation leads to several changes within the muscle compartment that may affect the mechanical interaction between adjacent muscles. Muscle fibers contract and, hence, the muscle belly shortens and expands radially. The former will result in a small change of the muscle relative position, whereas the expansion will increase the lateral tension of the connective tissue network.
In a recent study, we tested the hypothesis that the net effect of coactivation is an increase in the stiffness of the epimuscular pathways Figure 4which will facilitate force transmission between muscles. Effects of antagonist coactivation on mechanical interactions between synergistic muscles in the rat forelimb were assessed [ 36 ].
In contrast to the hypothesis, changes in force of the restrained muscle with length changes of its synergist were unaffected by antagonist coactivation. Testing intermuscular interaction with other combinations of active muscles e. Finally, it is also conceivable that the mechanical characteristics of the connective tissue system are different between muscle groups within an animal and across species.
The muscle-connective tissue architecture and composition of each synergistic group is different. Therefore, generalizing the current results to the whole musculoskeletal system should be done with caution. Although mechanical interactions between synergistic muscles have been shown in many animals e. In contrast to mammals, it has been reported that amphibians have a relatively poorly developed connective tissue network [ 37 ] and that insects contain very little connective tissue [ 3839 ].
Another aspect that should be taken into account is the scaling of muscle surface area to the 2nd power versus muscle volume to the 3rd power. This means that, for example, mice have a relatively larger epimysial surface to volume ratio than humans. To date, whether these variations between species lead also to different mechanical interactions remains unclear. The contradictory findings between the rat and cat studies are not fully understood and, thus, the responsible mechanisms requires further investigation.
Specifically, future studies should continue to test if the magnitude of intermuscular force transmission is dependent on the number of muscles that is simultaneously activated. Is the extent of force transmission between muscles the same throughout the body? This is another question that needs to be addressed. In conclusion, the importance of epimuscular myofascial pathways for muscle function during normal movements remains unclear.
Connective Tissue Function in Pathological Muscle-Tendon Conditions Besides a potential role for normal muscle function, epimuscular myofascial pathways may be important in pathological conditions of the musculoskeletal system. We found that when part of a muscle was normal the muscle as a whole generally developed the normal amount of tension and we guessed that some cell component other than the myofilament arrays served as a tension bypass through or around the damaged areas Ramsey and Street, unpublished.
This might promote healing. Enforcing music licenses is particularly problematic because retail musical recordings are readily available and copyright holders often literally must police unauthorized infringements in public places.How to find out what licenses and permits are needed to open a business
The sliding royalty scale can also lead to disputes when licensees believe they qualify for a lower rate than the licensing agency requests. Historically, this has sometimes provoked antagonism between music licensors like ASCAP and licensees like restaurants. As with music, software presents special problems for copyright holders.
Force Transmission between Synergistic Skeletal Muscles through Connective Tissue Linkages
By nature it is easily duplicated, but unlike music, software is typically not used in places open to the general public. This poses enormous challenges for enforcing software copyrights, as so-called pirated copies of software applications are believed to account for anywhere from 10 to 40 percent of the software installed on a typical business computer.
The problem is possibly more widespread in the consumer market. As with most copyright holders, software publishers have tried to closely restrict duplication and distribution of their products. As end user agreements, software licenses come in two forms: With a few exceptions, single user licenses allow only one installation of the program for use by one person.
Multi-user licenses usually specify a numeric range of permissible installations, such as up to 1, simultaneous users, and may restrict the physical space over which the software may be deployed, e. Some software installation programs include anti-piracy protections, but the same features that allow legal reuse of a program, for example, reinstalling the program after a system crash, tend to also permit illegal uses.
License infringements may occur at any of several stages in the software distribution process, and thus end users may not even know when they're using pirated copies. Some retailers and computer services install illegal copies of software on computer systems for sale as a means of attracting customers or boosting their profits by not paying licensing fees. There is also a thriving global black market for software in which illegal copies are transmitted for free or at minimal cost, with none of the proceeds returning to the publisher.
Businesses are generally liable for any illegal software on their computers, however, regardless of its origin. Individual software publishers also have methods of identifying piracy, such as through registration verification. When violations are found, large companies may be subject to fines in the hundreds of thousands of dollars.
Part of the solution to software licensing problems, some believe, is for businesses to implement sophisticated resource-tracking systems that monitor the installed base of programs—perhaps in real time—and identify when an unauthorized application has been installed. As of yet, there are few, if any, widely accepted tools for this, but it is an area of development likely to attract support from both software users and publishers. Aside from the contentious issues surrounding end-user software licenses, partner licensing is also a common practice within the software industry.
This occurs when one software developer licenses an application or, more often, an application component for use in conjunction with another developer's software.
Most major off-the-shelf software titles include some licensed components. For example, a graphics conversion filter in a word processing application may have been developed by an outside firm and purchased for either exclusive or nonexclusive use in the word processor.
One of the more common reasons is when two businesses claim rights to an invention. When one of them is ultimately determined to have the legitimate claim, the other must either license the patent from the its owner or give up the infringing portion of the work it has done on the competing version. The parties may choose to litigate it in court, or they may evaluate on their own the evidence and costs involved and settle with a licensing agreement.
A similar negotiation must occur when a company creates a patentable innovation that is based on someone else's patented product. In order for either company to use the innovation, the innovator must license its idea to the original patent holder, or the original holder must license its patent to the innovator.
As with other kinds of licensing agreements, patent licenses may be used for all sorts of business purposes. Sometimes the patent owner is a start-up inventor who needs an experienced partner to produce and market the invention.
For example, in the pharmaceutical and biotechnology industries, it is not uncommon for leading manufacturers to license some of their best-selling drugs from smaller firms, which in turn may derive a great deal of their revenue from licensing out their ideas rather than marketing them on their own.
At the same time, larger firms may wish to leverage their valuable patents in markets or applications that are outside their focus or expertise; for this they may choose smaller companies with unique credentials or other large firms with complementary market placement.
The semiconductor industry uses a special kind of engineering drawing known as a mask work to produce integrated circuits on semiconductor chips. Covered in the United States by the Semiconductor Chip Protection Act ofmask works enjoy somewhat of a cross between copyright and patent protection.
When they are registered they enjoy extended protection from duplication and competition. Walt Disney Company licenses a variety of its cartoon characters to Timex Corporation for use in character-themed watches bearing Timex's name. The food manufacturer ConAgra, Inc. Major universities license their athletic team logos to apparel producers in order to satisfy the strong public demand for college sports clothing and accessories and stimulate further interest in—and royalty revenue for—their athletic programs.
Each new character Disney creates in its animated movies is potentially licensed for dozens of merchandising applications such as toy figures, clothing, lunch boxes, and so forth. Depending on the circumstances, however, with licensed characters it may technically be a copyright that is licensed rather than a trademark. The process is similar, if less visible, with most kinds of trademark licensing, whether for the consumer market or the industrial market. Still, trademark licensors must also defend against unauthorized uses of their names and icons.
Particularly with apparel, they must contend with an extensive cottage industry of unlicensed apparel producers that use famous names and images to sell caps, T-shirts, and the like, often at a discount to "official" merchandise. Indeed, in some cases, as was true of Anheuser-Busch, it was out of defense against such abuses that companies first hit upon the idea of licensing their names and logos for profit.
Because trademark licensing builds on an already strong brand identity, once obvious precautions are taken against abuses it can represent one of the lowest-risk forms of licensing for both licensor and licensee alike.