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28 11. DE ARACHNIDIS<br />
3. MORPHOLOGY: EXTERNAL APPEARANCE 29<br />
work at Munich has shown that the organs are strictly mechanoreceptors,<br />
and that the minute deformations of the membrane, on<br />
which they depend, may be produced by:<br />
(i) vibration of the substrate;<br />
( ii) air-borne sounds;<br />
(iii) movements of parts of the body;<br />
(iv) the motion of walking.<br />
FIG. 5. Lyriform organ in section.<br />
"slit sense organs" they occur on the carapace and on the sternum and<br />
on the pedipalpi and legs, especially near the joints. Earth and Seyfarth<br />
( 1972) estimate as many as 3,000 on the surface of a single spider.<br />
During a long period of uncertainty several functions have been<br />
attributed to these objects. They have been described as auditory<br />
organs, chemoreceptors, detectors of heat and of atmospheric moisture.<br />
Pringle ( 1955), developing the idea that they are similar to the campaniform<br />
organs of insects, described them as modifications of the<br />
cuticle near the limb joints, with nerves or "sense cell processes"<br />
attached to the centre of a thin membrane. Thus he introduced the<br />
conception of mechanoreceptors, affecting orientation. Working with<br />
amputated legs from scorpions and Amblypygi, he attached to the<br />
bases of the nerves platinum electrodes connected to an amplifier and<br />
an oscilloscope. He found that pressure on the limb produced large<br />
impulses, and concluded that the lyriform organs supply a kinaesthetic<br />
sense, comparable to the muscle-tendon senses of vertebrates. Such a<br />
sense would play an important part in the phenomenon of Netzstarrheitstaxis<br />
of Holzapfel ( 1933).<br />
Similar results were obtained by Edgar (1963), using the legs of six<br />
species of Opiliones.<br />
Earth and Seyfarth ( 1972) have obtained striking results in favour of<br />
the orientation theory. In their apparatus a live spider, Cupiennius salei,<br />
was attracted by the buzzing of a bluebottle at distances ofup to 30 cm.<br />
On reaching the fly the spider was chased away and the fly removed.<br />
In normal circumstances and even with its eyes covered, the spider<br />
soon returned to the site of the insect. By immobilizing the lyriform<br />
organs on the femur or tibia the authors proved that these organs<br />
controlled kinetic orientation, a term that implies the determinaton of<br />
an animal's direction by its previous movements. Continuation of this<br />
(i) (ii) (iii)<br />
FIG. 6. Tarsal organs of spiders: (i) Myro; (ii) Bomis; (iii) Synaema.<br />
The tarsal organs (Fig. 6) are chemoreceptors and their use has<br />
been satisfactorily demonstrated by Blumenthal ( 1935). These organs<br />
can be seen under the microscope as small round holes in the upper<br />
surface of the tarsus, leading into a depression at the bottom of which<br />
lies a small projection. They are used in the testing of drinking water<br />
and the examination of edible prey. If the tarsus of one of the anterior<br />
legs of a thirsty spider is touched with a drop of water, the spider moves<br />
forward and drinks; if one of the posterior legs be so touched, the spider<br />
turns round and drinks. If the tarsal organ is sealed these responses<br />
disappear, and if a drop of water touches any other joint of the leg, the<br />
spider merely moves away. Blumenthal also showed that the organ<br />
enabled a spider to distinguish between water, brine, sugar solution and<br />
quinine.<br />
There is abundant evidence that most <strong>Arachnida</strong> react to sounds,