Jet propulsion

Octopuses swim headfirst, with arms trailing behind

Jellyfish pulsate their bell for a type of jet locomotion

Scallops swim by clapping their two shells open and closed

Main article: Jet propulsion

Jet propulsion is a method of aquatic locomotion where animals fill a muscular cavity and squirt out
water to propel them in the opposite direction of the squirting water. Most organisms are equipped
with one of two designs for jet propulsion; they can draw water from the rear and expel it from the rear,
such as jellyfish, or draw water from front and expel it from the rear, such as salps. Filling up the cavity
causes an increase in both the mass and drag of the animal. Because of the expanse of the contracting
cavity, the animal's velocity fluctuates as it moves through the water, accelerating while expelling water
and decelerating while vacuuming water. Even though these fluctuations in drag and mass can be
ignored if the frequency of the jet-propulsion cycles is high enough, jet-propulsion is a relatively
inefficient method of aquatic locomotion.

All cephalopods can move by jet propulsion, but this is a very energy-consuming way to travel compared
to the tail propulsion used by fish.[10] The relative efficiency of jet propulsion decreases further as
animal size increases. Since the Paleozoic, as competition with fish produced an environment where
efficient motion was crucial to survival, jet propulsion has taken a back role, with fins and tentacles used
to maintain a steady velocity.[11] The stop-start motion provided by the jets, however, continues to be
useful for providing bursts of high speed - not least when capturing prey or avoiding predators.[11]
Indeed, it makes cephalopods the fastest marine invertebrates,[12]:Preface and they can out accelerate
most fish.[13] Oxygenated water is taken into the mantle cavity to the gills and through muscular
contraction of this cavity, the spent water is expelled through the hyponome, created by a fold in the
mantle. Motion of the cephalopods is usually backward as water is forced out anteriorly through the
hyponome, but direction can be controlled somewhat by pointing it in different directions.[14] Most
cephalopods float (i.e. are neutrally buoyant), so do not need to swim to remain afloat.[10] Squid swim
more slowly than fish, but use more power to generate their speed. The loss in efficiency is due to the
amount of water the squid can accelerate out of its mantle cavity.[15]

Jellyfish use a one-way water cavity design which generates a phase of continuous cycles of jet-
propulsion followed by a rest phase. The Froude efficiency is about 0.09, which indicates a very costly
method of locomotion. The metabolic cost of transport for jellyfish is high when compared to a fish of
equal mass.
Other jet-propelled animals have similar problems in efficiency. Scallops, which use a similar design to
jellyfish, swim by quickly opening and closing their shells, which draws in water and expels it from all
sides. This locomotion is used as a means to escape predators such as starfish. Afterwards, the shell acts
as a hydrofoil to counteract the scallop's tendency to sink. The Froude efficiency is low for this type of
movement, about 0.3, which is why it's used as an emergency escape mechanism from predators.
However, the amount of work the scallop has to do is mitigated by the elastic hinge that connects the
two shells of the bivalve. Squids swim by drawing water into their mantle cavity and expelling it through
their siphon. The Froude efficiency of their jet-propulsion system is around 0.29, which is much lower
than a fish of the same mass.

Much of the work done by scallop muscles to close its shell is stored as elastic energy in abductin tissue,
which acts as a spring to open the shell. The elasticity causes the work done against the water to be low
because of the large openings the water has to enter and the small openings the water has to leave. The
inertial work of scallop jet-propulsion is also low. Because of the low inertial work, the energy savings
created by the elastic tissue is so small that it's negligible. Medusae can also use their elastic mesoglea
to enlarge their bell. Their mantle contains a layer of muscle sandwiched between elastic fibers. The
muscle fibers run around the bell circumferentially while the elastic fibers run through the muscle and
along the sides of the bell to prevent lengthening. After making a single contraction, the bell vibrates
passively at the resonant frequency to refill the bell. However, in contrast with scallops, the inertial work
is similar to the hydrodynamic work due to how medusas expel water - through a large opening at low
velocity. Because of this, the negative pressure created by the vibrating cavity is lower than the positive
pressure of the jet, meaning that inertial work of the mantle is small. Thus, jet-propulsion is shown as an
inefficient swimming technique.[15]

https://www.britannica.com/topic/locomotion/Saltation