Lecture 6 Fluid for skeletons
VorticesWhen I draw a canoe paddle through still water on a
lake, I see small vortices trail off the paddle edges. Coffee
stirring uses vortices to mix cream. Smokers blow smoke rings and
cetaceans blow bubble rings. There are crickets that communicate
with air vortices. This picture from the web is of a vortex forming
on the upstream side of a tidal turbine*. These rings may travel
through the fluid or stay fixed (e.g., over a drain). Fluids (both
water and air) exhibit flow and a vortex is simply flow that
spins.Toroid: doughnut-shaped object, e.g., O ring.
*the hole in the centre is ~15 cm in
diametersongofthepaddleRemember that air is a fluid just as
water.
Bio 325 Lecture 91Fluids are difficult to compress/effectively
incompressible, a fact which allows for translocation of forces by
hydrostatic skeletons (worms).This same fluid property of
incompressibility allows for changes of body shape hydraulically
when fluids are actually displaced (tube feet).And if fluid
incompressibility gets together with an opening its squeezing
results in expulsion: jet propulsion (squids).
Jet propulsion in plants: seed dispersal by detonation: sphagnum
spores as an example.
Assigned reading re sphagnum spores:Van Leeuwen J.L. 2010.
Launched at 36,000g. Science 329: 395-396.
Sphagnum or peat mossGenus of roughly 150-350 mosses in the
Phylum Bryophyta.Sphagnum grows in a thick carpet that can hold
15-20 times the moss's dry weight in water. Sphagnum creates wet,
acidic, anoxic conditions wherever it grows. conditions ideal for
the moss and inhospitable to competing species (modified from
Wikki). As a bryophyte, Sphagnum moss lacks a vascular system
(xylem, phloem), making it incapable of growing very tall above the
ground. Shortness makes difficulties for spore dispersal.
Most mosses disperse their spores by turbulent wind. Spore
capsules on 1-cm stalks cant reach boundary layer (10 cm up).Solved
by an air-gun mechanism ofexplosive discharge.Van Leeuwen 2010
Northwest of Upsala ON, Trans Canada HwyField Site Junior
WoodsyBlack spruce sphagnum bog4
Hyaline cells of Sphagnum showing pores that absorb water.
Spore capsules of Sphagnum, some already discharged are
cylindrical smaller, no cap; the more swollen have yet to dry
out.Epidermis of capsule dries in sun and shrinks, so capsule
volume decreases increasing internal air pressure ready for spore
discharge.
S. WhiteheadSpore must reach breezes that begin 10-15 cm above
the ground. Sphagnum shoots only ~1 cm, so spores have to be
propelled through a significant region of still air, a boundary
layer. Moreover, a spore capable of being carried by the wind must
also be subject to significant drag [keeping it aloft], meaning
that the ballistic launch of a [single] Sphagnum spore from the
ground would be incapable of propelling [it] to [the] necessary
height.6
Propelling small spores vertically is difficult because the low
terminal velocity that keeps them aloft also means they rapidly
decelerate in still air.
Definition of terminal velocity: when the sum of the drag force
and buoyancy equals the downward force of gravity: net force on an
object is zero and the object has zero acceleration.
J L van Leeuwen Science 2010;329:395-396Published by
AAASInitially spherical (A), the capsule containing spores is dried
by the sun, becoming (B) cylindrical at reduced volume and with
elevated internal pressure; the lid gives way and pressure/force
launches the spore mass, creating a toroid vortex ring. The jet of
spores and air rolls up into a turbulent ring vortex that carries
spores up to 15 to 20 cm. Without forming a vortex ring, which
keeps the spores clumped together the moss's spores would disperse
and fall uselessly to the ground...[from livescience.com] Quoting
Whitaker.
The vortex ring (toroid) generated in this context is a single
vortex, unlike the situation in jellyfish locomotion where vortices
follow each other in a series-- starting vortex alternating with
stopping vortex.The question I addressed in lecture: does the
expanding rising vortex ring, by its clockwise rotation adjacent to
the spore mass and the entrainment this implies, somehow contribute
to how high the spore mass rises? Can the vortex contribute to the
height gained by the spore
cluster?http://www.abc.net.au/science/articles/2010/07/23/2962175.htmSource:Mill
P.J. & Pickard R.S. 1975. Jet-propulsion in anisopteran
dragonfly larvae. J. comp. Physiol. 97(4): 329-338Some aquatic
insects use jetting to escape and to breathe.
Dragonfly nymphal/larval stageVentilation and Locomotion needs
satisfied by the same system. Abdomen a tagma telescoping for
changes in body volume.
Some dragonfly immatures (Order Odonata) jet water out of their
rectum (posterior gut chamber) which is also a respiratory chamber.
The insect uses a nozzle (see tapered terminal sclerites) to
increase momentum (mass X velocity) and to aim the jet flow just as
a squid siphon does. Telescoping in and out of abdominal segments
changes abdomen volume and so haemocoel pressure which powers water
intake and outflow; filaments filled with tracheae* project into
rectum lumen for gas exchange.nozzle*Gas exchange in insects via
tracheal system of air tubes.11Jetting used to launch at prey or
escape when disturbed or in water without a foothold.Jet propulsion
in insects is limited to dragonfly larvae; note this Mill &
Pickard paper written with comparison in mind, e.g., to squids
(Cephalopoda).
Cycles of nymph jetting repeated at 2.2 per s for sustained
progress; see Mill & Pickards longitudinal and oblique muscles
diagram of 9th and 10th abdominal segments.The effectiveness of the
jet-propulsion mechanism is largely dependent upon a) velocity and
mass of water ejected from the respiratory chamber b) the mass of
the whole animal and c) magnitude of induced drag forces. cuticular
restoring forces.
Mill P.J., Pickard R.S. 1975. Jet-propulsion in anisopteran
dragonfly larvae. J. comp. Physiol. 97(4): 329-338.
A colony of zooids: at one end specialized as swimming
individuals called nectophores, also called swimming bells. They
jet seawater out of their subumbrellar openings, moving the colony
that trails behind on a fishing stem (stem transports nourishment
by a shared coelenteron).Kevin RaskoffNational GeographicNectophore
zooids in a hydrozoan Cnidarian13Tunicate: Urochordata
Sea squirts are small barrel-shaped creatures often living in
clusters. They have a tadpole larva dispersal phase with a
notochord, absent in adults. An incurrent siphon, see os above,
brings seawater into a slitted pharynx which filters food;
excurrent exits at ats. These two siphons have been adapted in some
species for locomotion: see salps.Barrington, E.J.W. 1965. The
Biology of Hemichordata and Protochordata. Oliver & Boyd,
Edinburgh, London.
Scanned from Barrington, p. 84; os inhalant siphon, ats exhalant
siphon. In B ic is the direction of the incurrent which moves out
of the pharynx lumen through the pharynx slits into an atrial
cavity, thence to the ats. The current is created by beating cilia.
Sheets of mucus on the pharynx walls trap the diatoms and other
tiny organisms that are then concentrated and passed on down the
gut. This sessile filter feeder, the species is Clavelina
lepadiformis, is an example of a more typical tunicate. Compare
with the salps.14
Peter J. BryantA propulsive jet for locomotion is created by
rhythmic compression of muscle bands encircling the barrel-shaped
body. Fluid enters the anterior oral siphon to fill the mostly
hollow body of the salp. ...oral lips close and circular muscle
bands contract, decreasing the volume of the jet chamber so that
fluid is accelerated out of the posterior atrial siphon.
[antagonists?] ...unique in possessing incurrent and excurrent
siphons on opposite ends of the body allowing for unidirectional
flow and reverse swimming during escape (Sutherland et al.
2010).Some tunicates called salps are Pelagic: living in the open
ocean and swimming by ejecting seawater. See: Sutherland K.R.,
Madin L.P. 2010. Comparative jet wake structure and swimming
performance of salps. J. exp. Biol. 213: 2967-2975.Pelagic: any
water in a sea or lake that is neither close to the bottom or the
shore (Wikki) open ocean15
Sutherland Fig. 3A propulsive jet for locomotion is created by
rhythmic compression of muscle bands encircling the barrel-shaped
body. Fluid enters the anterior oral siphon to fill the mostly
hollow body of the salp. Oral lips close and circular muscles
contract, decreasing chamber volume so that fluid is accelerated
out of the posterior atrial siphon. [antagonists?]...unique in
possessing incurrent and excurrent siphons on opposite ends of the
body allowing for unidirectional flow and reverse swimming during
escape.16Assigned reading: Sutherland K.R., Madin L.P. 2010.
Comparative jet wake structure and swimming performance of salps.
J. exp. Biol. 213: 2967-2975.
Kenneth KoppSalp chains: individuals (zooids) strung together in
coloniesColonial tunicatesI wonder if vortex-assisted locomotion
(vortex-ring propulsion) is not universal in jetting animals?Are
there any fish that employ jet propulsion? Flounders (flatfish) use
their offside operculum. See Brainerd E.L., Page B.N., Fish F.E.
1997. Opercular jetting during fast-starts by flatfishes. J. exp.
Biol. 200: 1179-1188.17Jellyfish Jetting locomotionAssigned
reading: Dabiri J. O., Colin S.P., Costello J.H., Gharib, M. 2005.
Flow patterns generated by oblate medusan jellyfish: field
measurements and laboratory analyses. Journal of experimental
Biology 208: 1257- in which they demonstrate the stopping vortex
ring which contributes to medusa swimming.
Yong, Ed 2013. Why a jellyfish is the oceans most efficient
swimmer. Nature doi:10.1038/nature.2013.13895
JELLYFISH FORM AND FUNCTION Website by John H. Costello &
Sean P. Colin, Roger Williams University. See this website for
information about jellyfish swimming form from specialists:
>fox.rwu.edu/jellies/index.html