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Topic: Halteres

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	Abstract Nalbach: "The halteres of the blowfly Calliphora. 1." (Site not responding. Last check: 2007-10-10)
		Abstract Nalbach: "The halteres of the blowfly Calliphora.
		The physics of the halteres are described in terms of a general formula for the force acting onto the endknob of the moving haltere during rotations and linear accelerations of the fly (Eq.
		On the basis of the experimentally determined kinematics of the haltere, the primary forces and the forces dependent on angular velocity and on angular acceleration are calculated (Figs.
	www.kyb.tuebingen.mpg.de /misc/abstr93/nalbac_1.html (237 words)

	Haltere Afferents Provide Direct, Electrotonic Input to a Steering Motor Neuron in the Blowfly, Calliphora -- ...
		Halteres are sophisticated equilibrium organs of flies that function to detect angular rotations of the body during flight.
		Both the size of the EPSP and the activity pattern in the haltere nerve were sensitive to the precise alignment of the plane in which the haltere was oscillated, indicating that the campaniform sensilla were quite sensitive to the direction of cuticular strain.
		The delay between haltere nerve stimulation and the mnb1 EPSP is comparable to the delay between the extracellularly recorded spike in the haltere nerve and the unitary EPSP in mnb1 in response to mechanical stimulation of a single campaniform in dF2.
	www.jneurosci.org /cgi/content/full/16/16/5225 (6975 words)

	Halteres (Site not responding. Last check: 2007-10-10)
		Halteres, (singular halter or haltere) from the Greek word for dumbbells, are small knobbed structures homologous to wings and flapped to maintain stability when flying.
		In ancient Greek sports, halteres were used as lifting weights, and also as weights in their form of the long jump.
		Their long jump was probably a set of three jumps, and halteres were held in both hands to allow an athlete to jump a greater distance.
	publicliterature.org /en/wikipedia/h/ha/halteres.html (177 words)

	Halteres
		In ancient Greek sports, halteres were used as lifting weights, and also as weights in their version of the long jump, which was probably a set of three jumps.
		Halteres were held in both hands to allow an athlete to jump a greater distance; they may have been dropped after the first or second jump.
		Halteres were made of stone or metal, and weighed between two and nine kilograms.
	www.mlahanas.de /Greeks/LX/Halteres.html (163 words)

	A long jump - Letters - Letter to the Editor Natural History - Find Articles
		Halteres may have been used at times for the standing long jump, but probably not for the competitive event.
		Biologists have adopted the term "halteres" to refer to the rear vestigial "wings," or balancers, in dipterans (two-winged flies, mosquitoes, gnats, and so on).
		Curiously, in the fruit fly, a single gene mutation is capable of making the halteres revert to a second set of wings, thus anatomically removing the mutated flies from the order Diptera.
	www.findarticles.com /p/articles/mi_m1134/is_5_112/ai_102275143 (477 words)

	04.09.98 - UC Berkeley scientists find key to fly's maneuverability and the speed with which they elude fly swatters
		The halteres then relay signals to the wing muscles to alter their stroke or angle of attack.
		The halteres, beating out of sync with the forewings, are the key to the fly's aerodynamic prowess.
		The halteres, just one nerve cell away from the motor neurons of the wing, are designed to react quickly - reflexively - to yaw, pitch and roll in the fly.
	www.berkeley.edu /news/media/releases/98legacy/04_09_98a.html (1276 words)

	Neurobehavior: Module 3
		If it has more than 1 pair then it must not have halteres, becuase halteres are only found in insects that have a reduced 2nd pair of wings that have evolved into sense organs that detect changes in the insect's orientation = halteres.
		Halteres are sensory structures that detect a change in the orientation of an insect in flight.
		When the halteres detect a change they trigger motor neurons to correct for the change and thus stabilize the insect while in flight.
	www.lifesci.ucsb.edu /~mcdougal/neurobehavior/modules_homework/mod3.html (924 words)

	Why a fly can fly like a fly
		The halteres tend to keep the insect flying in the same direction, so when it has to make a sudden turn, the halteres have to respond as well as the flight muscles.
		So rather than ‘switching off one’s gyroscope’;, the halteres are appropriately fine-tuned or ‘tweaked’ for the required manoeuvre in response to visual cues, a fraction of a second before they in turn pass the information on to the main flight muscles.
		Experimental evidence indicates that halteres are part of a highly sophisticated, and immensely complex, flight system which continues to baffle and amaze all who study it.
	www.answersingenesis.org /tj/v12/i3/fly.asp (1065 words)

	Michael Dickinson and Claire Balint Q&A
		In the case of the fly, the beating halteres are subject to a force whenever the body of the fly rotates during flight.
		Tiny sensors at the base of the haltere detect these forces which tend to push and pull the globular end of the haltere in and out of its narrow stroke plane.
		The haltere sensors are connected to wing steering muscles that change the way the wing moves in order to counter-act the imposed rotation.
	www.pbs.org /safarchive/3_ask/archive/qna/3294_bees.html (1853 words)

	Exponent-News & Press Releases
		The jump distance for the nominal simulation (no halteres used) was 2.35 m, while the jump distance for the simulation that used halteres was 2.74 m, an improvement of 39 cm.
		Analysis of the simulation mechanics showed that 12 cm of this improvement was due to the greater horizontal position of the center of gravity at take-off from extending the halteres ahead of the body.
		Releasing the halteres during flight increased the jump performance an additional 15 cm, for a total jump distance of 2.89 m.
	www.exponent.com /about/news/archives/2005/10/24/20.42.14 (838 words)

	Halteres - Wikipedia, the free encyclopedia
		Crane fly, with a pair of halteres visible below the wings as appendages, about as long as the animal's antennae, with knobs on the end
		Halteres are homologous to, and evolved from, insect wings.
		In the Strepsiptera the forewings changed into halteres, while in the Diptera (flies, mosquitoes and gnats) the hindwings evolved into halteres.
	en.wikipedia.org /wiki/Halteres (252 words)

	Physics Today January 2003
		As the jumper prepares to land, the halteres are swung down and backward so that, on landing, the feet of the weighted jumper are more forward of the center of mass than they are for modern, unweighted athletes.
		The greater takeoff speed achieved by loaded jumpers led to increases in distances beyond those generated by using the halteres to shift the position of the feet relative to the center of mass.
		Written testimonies stipulate that the foot had to land "properly," and that the halteres were an aid to proper landing.
	www.physicstoday.org /vol-56/iss-1/p15.html (801 words)

	04.15.98 - Learning How Flies Fly
		The halteres relay signals to the wing muscles to alter their stroke or angle.
		Though these steering muscles – 11 of them in the house fly, analogous to the 17 steering muscles attached to the fly’s forewing – evidently are no longer important in generating aerodynamic forces, Dickinson had a hunch they might be the missing connection between the visual system and the flight muscles.
		The halteres, just one nerve cell away from the motor neurons of the wing, are designed to react quickly – reflexively – to yaw, pitch and roll in the fly.
	www.berkeley.edu /news/berkeleyan/1998/0415/flies.html (1125 words)

	Arthropods of Economic Importance - Agromyzidae of the World > Higher groups
		They are generally characterized by the single pair of membranous wings and the halteres, which are homologous to the hind wings in other groups of winged insects.
		Although the halteres can be viewed as atrophied hind wings, they have an important function for the fly's flight.
		The halteres, during flight operating as oscillating pendula, serve as sensual organ.
	ip30.eti.uva.nl /bis/agromyzidae.php?menuentry=groepen (976 words)

	The Ancient Olympics
		To increase jumping distance, the athlete held one of these telephone receiver or dumbbell shaped weights in each hand, ran and jumped, swinging the weights, and released the halteres behind him at the end of his jump.
		Halteres weighed between 1.6 to 4.6 kilograms, or 3.5 to 10.1 pounds.
		Pausanias describes the halteres as "half of a circle, not an exact circle but elliptical, and made so that the fingers pass through as they do through the handle of a shield," Description of Greece, 5.26.3.
	ablemedia.com /ctcweb/consortium/ancientolympics13.html (499 words)

	EducationGuardian.co.uk \| Special Reports \| How the Greeks jumped to it
		Halteres were swung backwards and forwards by the competitors before takeoff, then thrust forwards during the first part of the flight, and finally back just before landing.
		If, just before landing, athletes throw their halteres behind them, they gain more edge, because, as the halteres fly backwards, their feet move an few centimetres forwards.
		Halteres that weighed too little were of no help, and any extra advantage disappeared at around 10kg.
	education.guardian.co.uk /museums/story/0,11727,839652,00.html (382 words)

	Dickinson Lab - Visual-Mechanosensory Fusion in Flight Equilibrium
		The main way that flies maintain stable flight is through sensory feedback from their visual system and their mechanosensory haltere system.
		The halteres are modifications of the hindwings (recall that dragonflies have four wings, while "true" flies are defined by having only two) into complex sensors that detect Coriolis forces proportional to the angular velocity of the fly.
		Thus, if a freely flying fly were to be rotated by an external force such as wind, its visual system would be capable of detecting this rotation, but the haltere system would also be stimulated, and is capable of initiating a compensatory response even in the absence of visual feedback.
	www.its.caltech.edu /~flight/research_visualmechanofusion.html (160 words)

	The Ancient Olympics- Jumping (Site not responding. Last check: 2007-10-10)
		Jumping was directly related to the skills that a soldier must possess in war.
		The jumper held the halteres in front of him during
		halteres made it extremely difficult for the athletes to remain co-ordinated and so flute music was played during the
	www.personal.psu.edu /students/s/u/sua132/Jumping.htm (122 words)

	The Initiation and Control of Rapid Flight Maneuvers in Fruit Flies -- Dickinson 45 (2): 274 -- Integrative and ...
		Halteres are the tiny club-shaped hindwings characteristic of
		Descending visual input is thought to converge with mechanosensory input from wing and haltere on motor neurons of steering muscles such as b1.
		Pringle, J. The gyroscopic mechanism of the halteres of Diptera.
	icb.oxfordjournals.org /cgi/content/full/45/2/274 (4073 words)

	Summation of visual and mechanosensory feedback in Drosophila flight control -- Sherman and Dickinson 207 (1): 133 -- ...
		of the linear velocity of the haltere and the angular velocity of
		Input from the halteres and visual system are combined in a weighted sum by the flight control system.
		The weighting of the visual feedback is dependent on the presence of haltere input but appears to be constant over a wide range of haltere stimulation.
	jeb.biologists.org /cgi/content/full/207/1/133 (5805 words)

	ecowatch_diptera (Site not responding. Last check: 2007-10-10)
		Although little is known of the Australian species, the larvae of bee flies are believed to parasitise the larvae of other insects and may prey on the eggs in egg-masses of grasshoppers and locusts.
		The base of halteres are flexible and when they are moved a fly or mosquito is able to control its flight.
		As the haltere bends at the base, a fly or mosquito can change flight speed or direction making them more manoeuvrable compared to many other flying insects.
	www.ento.csiro.au /Ecowatch/Diptera/diptera.htm (520 words)

	eNaturalist: Giant Mosquito? (quick read) (Site not responding. Last check: 2007-10-10)
		The halteres vibrate as the animal navigates and allow a fly to monitor changes in direction and attitude.
		This activity should make it clearer exactly what a haltere looks like and we hope will also firmly establish one of a fly’s key anatomical characteristics in their minds.
		Incidentally, the reason we know that flies can’t fly without their halteres is that researchers captured some live flies and removed their halteres as an experiment and found that each fly was rendered totally incapable of flight.
	www.enaturalist.org /unit/147/qr (1427 words)

	The Citizen Scientist
		When the crane fly switches on its powerful wing motor, the halteres whip back and forth in time with the wings.
		Inertia tends to keep the vibrating halteres from moving away from their former position when the fly makes a turn.
		This causes the stalk of the haltere to activate sensors behind the wing that tell the fly in which direction and exactly how far it has turned.
	www.sas.org /tcs/weeklyIssues_2005/2005-04-22/mimsci/index.html (641 words)

	FLIES (Site not responding. Last check: 2007-10-10)
		Diptera is the name given to true flies, the name means two wings and it is this feature which differentiates them from other types of fly such as mayflies and alder flies which have four wings.
		The function of these halteres is to maintain stable flight by signaling to the brain the flies spatial orientation.
		This is achieved by small groups of campaniform sensilla which detect mechanical stress imposed upon the cuticle at the base of the vibrating halteres.
	www.roberth.u-net.com /Diptera.htm (630 words)

	The Official Site of the Houston Texans (Site not responding. Last check: 2007-10-10)
		Historians tell us the halteres was a piece of athletic equipment used during competition by long jumpers in Ancient Greece.
		Athletes would attempt to simultaneously jump and throw the halteres forward hoping it would propel them a greater distance.
		The halteres was also used to perform strengthening exercises to help athletes prepare for athletic competitions.
	www.houstontexans.com /news/news_detail.php?PRKey=250 (968 words)

	Ancient Olympians: Weighted Down to Win (Site not responding. Last check: 2007-10-10)
		The halteres could also help the athletes while they were mid-bound, even though the trajectory of a jumper's center of mass is determined at the instant of launch.
		Furthermore, by enabling competitors to bring their upper-body muscles into play during launch, halteres would have permitted jumpers to propel themselves off the ground with more initial force—at least enough to compensate for the extra weight they bore.
		Even if halteres were permitted today, he noted, they wouldn't necessarily propel long-jumpers farther.
	news.nationalgeographic.com /news/2002/11/1114_021114_GreekAthletes_2.html (582 words)

	A comparison of visual and haltere-mediated equilibrium reflexes in the fruit fly Drosophila melanogaster -- Sherman ...
		Halteres are small evolutionarily modified hind wings (Fig.
		Halteres encode faster rotations whereas the visual system is responsive to slower rotations along all three functional axes.
		Strausfeld, N. and Seyan, H. Convergence of visual, haltere, and prosternal inputs at neck motor neurons of Calliphora erythrocephala.
	jeb.biologists.org /cgi/content/full/206/2/295 (4455 words)

	Sep 95 - Bayfront Byline Bug Walk (Site not responding. Last check: 2007-10-10)
		Behind each wing sticking out from the thorax (like a pin stuck in a pin cushion) is a greatly modified wing called a haltere, a slender stalk topped with a knob.
		Halteres are sense organs which are essential for stable flight.
		In flight, the halteres move up while the wings are moving down and down while the wings are moving up.
	cassfos02.ucsd.edu /personal/ron/CVNC/byline/bugs_95sep.html (397 words)

	True Flies - Order Diptera
		These forewings are located on the mesothorax, and since flies must rely only on this pair of wings for flight the mesothorax is slightly enlarged to contain the flight muscles.
		Special organs known as halteres are located behind the forewings.
		Flies are, as their common name suggests, wonderful fliers, many species being able to fly forwards and backwards at remarkable speeds and even hover in midair.
	www.angelfire.com /mo2/animals1/arthropod/diptera.html (696 words)

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