A muscular hydrostat is a biological structure found in animals.

It is used to manipulate items

(including food) or to move its host about and consists mainly of muscles with no skeletal support. It
performs its hydraulic movement without fluid in a separate compartment, as in a hydrostatic
skeleton.

A muscular hydrostat, like a hydrostatic skeleton, relies on the fact that water is effectively
incompressible at physiological pressures. In contrast to a hydrostatic skeleton, where muscle
surrounds a fluid-filled cavity, a muscular hydrostat is composed mainly of muscle tissue. Since muscle
tissue itself is mainly made of water and is also effectively incompressible, similar principles apply.

Muscular anatomy

Closeup of the trunk of an Asian elephant

Muscles provide the force to move a muscular hydrostat. Since muscles are only able to produce force
by contracting and becoming shorter, different groups of muscles have to work against each other,
with one group relaxing and lengthening as the other group provides the force by contracting. Such
complementary muscle groups are termed antagonistic pairs.

The muscle fibers in a muscular hydrostat are oriented in three directions: parallel to the long axis,
perpendicular to the long axis, and wrapped obliquely around the long axis. [1][2]

The muscles parallel to the long axis are arranged in longitudinal bundles. The more peripherally these
are located, the more elaborate bending movements are possible. A more peripheral distribution is
found in tetrapod tongues, octopus arms, nautilus tentacles, and elephant trunks. Tongues that are
adapted for protrusion typically have centrally located longitudinal fibers. These are found in snake
tongues, many lizard tongues, and the mammalian anteaters.

The muscles perpendicular to the long axis may be arranged in a transverse, circular, or radial pattern.
A transverse arrangement involves sheets of muscle fibers running perpendicular to the long axis,
usually alternating between horizontal and vertical orientations. This arrangement is found in the
arms and tentacles of squid, octopuses, and in most mammalian tongues. A radial arrangement
involves fibers radiating out in all directions from the center of the organ. This is found in the tentacles
of the chambered nautilus and in the elephant proboscis (trunk). A circular arrangement has rings of
contractive fibers around the long axis. This is found in many mammalian and lizard tongues along
with squid tentacles.
Helical or oblique fibers around the long axis are generally present in two layers with opposite chirality
and wrap around the central core of musculature.

Mechanism of operation

In a muscular hydrostat, the musculature itself both creates movement and provides skeletal support
for that movement. It can provide this support because it is composed primarily of an incompressible
“liquid" and is thus constant in volume. The most important biomechanical feature of a muscular
hydrostat is its constant volume. Muscle is composed primarily of an aqueous liquid that is essentially
incompressible at physiological pressures. In a muscular hydrostat or any other structure of constant
volume, a decrease in one dimension will cause a compensatory increase in at least one other
dimension.[3] The mechanisms of elongation, bending and torsion in muscular hydrostats all depend on
constancy of volume to effect shape changes in the absence of stiff skeletal attachments. [4] Since
muscular hydrostats are under constant volume when the diameter increases or decreases, the length
must also decrease or increase, respectively. When looking at a cylinder the volume is: V=πr²l. When
the radius is differentiated with respect to the length: dr/dl=-r/(2l). From this, if a diameter decreases
by 25%, the length will increase by approximately 80% which may produce a large amount of force
depending on what the animal is trying to do.[5]

Elongation and shortening

Arms and tentacles of the squid Abralia veranyi

Elongation in hydrostats is caused by the contraction of transverse or helical musculature

arrangements. Given the constant volume of muscular hydrostats, these contractions cause an
elongation of the longitudinal muscles. Change in length is proportional to the square of the decrease
in diameter.[3] Therefore, contractions of muscles perpendicular to the long axis will cause a decrease
in diameter while keeping a constant volume will elongate the organ length-wise. Shortening, on the
other hand, can be caused by contraction of the muscles parallel to the long axis resulting in the organ
increasing in diameter as well as shortening in length.

The muscles used in elongation and shortening maintain support through the constant volume
principle and their antagonistic relationships with each other. These mechanisms are seen often in
prey capture of shovelnose frogs and chameleons, as well as in the human tongue and many other
examples. In some frogs, the tongue elongates up to 180% of its resting length. [6] Extra-oral tongues
show higher length/width ratios than intra-oral tongues, allowing for a greater increase in length
(more than 100% of resting length, as compared to intra-oral tongues at only about 50% of resting
length increase). Greater elongation lengths trade off with the force produced by the organ; as the
length/width ratio is increased elongation increases while force is decreased. [1] Squids have been
shown to use muscular hydrostat elongation in prey capture and feeding as well.[7]

Bending
The bending of a muscular hydrostat can occur in two ways, both of which require the use of
antagonistic muscles.[1] The unilateral contraction of a longitudinal muscle will produce little or no
bending and will serve to increase the diameter of the muscular hydrostat because of the constant
volume principle that must be met. To bend the hydrostat structure, the unilateral contraction of
longitudinal muscle must be accompanied by contractile activity of transverse, radial, or circular
muscles to maintain a constant diameter. Bending of a muscular hydrostat can also occur by the
contraction of transverse, radial, or circular muscles which decreases the diameter. Bending is
produced by longitudinal muscle activity which maintains a constant length on one side of the
structure.

The bending of a muscular hydrostat is particularly important in animal tongues. This motion provides
the mechanism by which a snake flicks the air with its tongue to sense its surroundings, and it is also
responsible for the complexities of human speech.[2]

Stiffening

The stiffening of a muscular hydrostat is accomplished by the muscle or connective tissue of the
hydrostat resisting dimensional changes.[3]

Torsion

Torsion is the twisting of a muscular hydrostat along its long axis and is produced by a helical or
oblique arrangement of musculature[3] which have varying direction. For a counter-clockwise torsion it
is necessary for a right-hand helix to contract. Contraction of a left-hand helix causes clockwise
torsion. The simultaneous contraction of both right and left-hand helixes results in an increase in
resistance to torsional forces. The oblique or helical muscle arrays in the muscular hydrostats are
located in the periphery of the structure, wrapping the inner core of musculature, and this peripheral
location provides a larger moment through which the torque is applied than a more central location.
The effect of helically arranged muscle fibers, which may also contribute to changes in length of a
muscular hydrostat, depends on fiber angle—the angle that the helical muscle fibers make with the
long axis of the structure.

The length of the helical fiber is at a minimum when the fiber angle equals 54°44 ′ and is at maximum
length when the fiber angle approaches 0° and 90°.[3] Summed up, this means that helically arranged
muscle fibers with a fiber angle greater than 54°44′ will create force for both torsion and elongation
while helically arranged muscle fibers with a fiber angle less than 54°44 ′ will create force for both
torsion and shortening.[8] The fiber angle of the oblique or helical muscle layers must increase during
shortening and decrease during lengthening. In addition to creating a torsional force, the oblique
muscle layers will therefore create a force for elongation that may aid the transverse musculature in
resisting longitudinal compression.

Examples
 West Indian manatee

 Whole bodies of many worms

 Feet of mollusks[9] (including arms and tentacles in cephalopods[1][7][10][11])

 Tongues of mammals and reptiles[1]

 Trunks of elephants[1]

 The snout of the West Indian manatee[12]

Technological applications

A group of engineers and biologists have collaborated[when?] to develop robotic arms that are able to
manipulate and handle various objects of different size, mass, surface texture and mechanical
properties. These robotic arms have many advantages over previous robotic arms that were not based
on muscular hydrostats.[13]

References

1.

 Kier, W. M.; Smith, K. K. (1985). "Tongues, tentacles and trunks: The biomechanics of movement in
muscular-hydrostats". Zoological Journal of the Linnean Society. 83 (4): 307–324. doi:10.1111/j.1096-
3642.1985.tb01178.x.

  Smith, Kathleen K.; William M. Kier (Jan–Feb 1989). "Trunks, tongues, and tentacles: Moving with
skeletons of muscle". American Scientist. 77 (1): 28–35.

  Kier, W. M. (1985). "The musculature of squid arms and tentacles: Ultrastructural evidence for
functional differences". Journal of Morphology. 185 (2): 223–239. doi:10.1002/jmor.1051850208.
PMID 30011972. S2CID 51631466.

  Wainwright, P. C.; Bennett, A. F. (1992). "The mechanism of tongue projection in chameleons: II.
Role of shape change in a muscular hydrostat". The Journal of Experimental Biology 168: 23–40.

  Alexander, R. McN. (2003). Principles of Animal Locomotion. Princeton, NJ: Princeton University
Press.

  Nishikawa, K. C.; Kier, W. M.; Smith, K. K. (1999). "Morphology and mechanics of tongue
movement in the African pig-nosed frog Hemisus marmoratum: a muscular hydrostatic model
http://jeb.biologists.org/content/202/7/771.short". The Journal of Experimental Biology 202: 771–
780.

  Kier, W. M. (1982). "The functional morphology of the musculature of squid (Loliginidae) arms
and tentacles". Journal of Morphology. 172 (2): 179–192. doi:10.1002/jmor.1051720205.
PMID 30103569. S2CID 51980244.

  Meyers, J. J.; O'Reilly, J. C.; Monroy, J. A.; Nishikawa, K. C. (2004). "Mechanism of tongue
protraction in microhylid frogs. The Journal of Experimental Biology 207: 21–31.

  Price, Rebecca (Dec 2003). "Columellar muscle of neogastropods: Muscle attachment and the
function of columellar folds". Biological Bulletin. 205 (3): 351–366. doi:10.2307/1543298.
JSTOR 1543298. PMID 14672989. S2CID 12277966.

  Matzner, H.; Gutfreund, Y.; Hochner, B. (2000). "Neuromuscular system of the flexible arm of the
octopus: Physiological characterization". Journal of Neurophysiology. 83 (3): 1315–1328.
doi:10.1152/jn.2000.83.3.1315. PMID 10712459. S2CID 14402766.

  Yekutieli, Y.; Sumbre, G.; Flash, T.; Hochner, B. (2002). "How to move with no rigid skeleton? The
octopus has the answers". Biologist (London, England). 49 (6): 250–254. PMID 12486300.

  Marshall, C. D.; Clark, L. A.; Reep, R. L. (1998). "The muscular hydrostat of the Florida manatee
(Trichechus manatus latirostris): A functional morphological model of perioral bristle use". Marine
Mammal Science. 14 (2): 290–303. doi:10.1111/j.1748-7692.1998.tb00717.x.

13.  Walker, I.D.; Dawson, D.M.; Flash, T.; Grasso, F.W.; Hanlon, R.T.; Hochner, B.; Kier, W.M.;
Pagano, C.C.; Rahn, C.D.; Zhang, Q.M. (2005). "Continuum robot arms inspired by
cephalopods". Proceedings of SPIE 5804: 303–314.

Categories:

 Animal anatomy

 Biomechanics

 This page was last edited on 16 April 2024