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Findings on the Electrograms of the Cicada's Heart (Cryptotympan a Japonensis Kato)

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Findings on the Electrograms of the Cicada's Heart (Cryptotympan a Japonensis Kato) FINDINGS ON THE ELECTROGRAMS OF THE CICADA'S HEART (CRYPTOTYMPAN A JAPONENSIS KATO) HIROSHI IRISAWA, AYA F. IRISAWA AND TETSUJI KADOTANI Department of Physiology, School of Medicine, Hiroshima University Several works on the neurogenic hearts of arthropoda by Carlson (1) and Crescitelli (2, 9) have already shown that their electrocardiograms are markedly different from these of vertebrates. One of the characteristics of the electro- grams of the arthropoda heart is that they show oscillatory changes of potentials which have been attributed to rhythmical discharges of the pacemakers (9, 5, 7). Some of the previous workers (4, 13) have shown that under normal con- ditions the electrogram exhibits simpler patterns. However, it seems that their work did not show anything about the pacemaker activities of the heart beat. The present paper deals with the experimental results of the localization of pacemakers in the cicada's heart. METHODS A schematic sketch of the cicada's heart (Cryptotymana japonensis Kato) is shown in figure 1. It consists of seven segments, each of which is stretched by a pair of alary muscles. The microscopic sturucture is illustrated in figure 2. The surface of heart is covered with several layers of cubic cells. Beneath these layers there is a layer of striated muscle fibers about 20ƒÊ in diameter which are loosely connected with each other unlike those of the mammalian myocardium. For electrical observations, two types of electrode were used. 1) A low- resistance microelectrode. originally reported by Tomita et al. (17), which con- sists of about a 5ƒÊ silver wire, covered with a glass tubing, with an outer dia- meter of 10-15ƒÊ. By means of this electrode the potential can be led off from much smaller areas, than those of previous workers who used such electrodes with tip diameters exceeding 100ƒÊ for the insect heart. The heart action po- tential was recorded by a cathode ray oscilloscope through a 4 stage CR-ampli- fier. Cicada was placed in Ringer solution, to which the indifferent electrode was connected. 2) In order to record the membrane potential of heart muscle fiber intracellular electrodes of the type employed by Nastuk and Hodgkin were used (14). They were filled with 3 M KCl solution by Tasaki's method and those with electrical resistances of about 20 Megohms were selected. For the recording a cathode follower preamplifier (954) with very small grid current Received for publication December 23, 1956. *入 沢 宏 ,入 沢 彩,角 谷 哲 司 150 ELECTROGRAMS OF THE CICADA'S HEART 151 was connected with the main amplifier. Both low-and high-resistance micro- electrodes were moved by a micromanipulator under the microscope. FIG.2.•@ Photomicrograph of cicada's heart. M: Striated muscle. Parallel fibers about20ƒÊ in diameter. A: Alary muscle attached to heart surface. V: Valves located at ostium. FIG.1.•@ Schematic sketch of cicada's heart. Roman numerals label each segment. EXPERIMENTAL RESULTS 1) Electrogram from the intact heart: In order to observe the heart tube, the abdomen was opened, and the muscle and digestive organs were removed. Figure3shows recordings of intricate surface action potentials led off with a low resistance electrode from various parts along the heart. At the anterior part of heart, the recording consists of several small and one large action potential. Usually this tendency was observed almost every- where on the heart tube. In this instance, however, diphasic action potentials were led off from the5th and 6th segment, while slow monophasic potential changes were observed at the7th segment. From these pictures, it can be seen that the electrograms of intact cicada's heart are so complex, that it seems to be very difficult to analyse them as a whole. 2) Electrogram from the isolated heart: As the intricacy of the electrogram described above, seemed to be mainly due to the alary muscles, the isolation of the heart tube per se from the alary muscles was tried. No remarkable changes of heart contraction were observed as a result of the removal of the alary muscles. Figure4illustrates the surface action potentials of the extirpated heart which are simpler than those in figure3. As would be expected almost all of the action potentials were triphasic, except two tracings from the anterior part of heart. At the neighbourhood of the7th segment, the initial deflection 152 H. IRISAWA ET AL. FIG.3.•@ Electrograms from the intact cicada's heart. Right: Schematic sketch of the cicada's heart. Left: Action po- tentials recorded from the place indicated. Up- ward deflection in elec- trograms is positive. Time maker: 0.3sec. FIG.4.•@ Surface action po- tentials from isolated cicada's heart. Recorded from indicat- ed locations. ELECTROGRAMS OF THE CICADA'S HEART 153 of tracing is negative. This negative sign localized in and around the 7th seg- ment. If the 7th segment is cutted off, this negative deflection would be obser- ved only at the 2nd segment. In the isolated heart, another very special finding was obtained from a particular part near the end of the 7th segment. As shown in the bottom tracing of figure 4, this was a very regular positive monophasic action potential with a different frequency from that of other parts of heart, and it seems that this potential localized only at the 7th segment and was not conducted. 3) The action potentials from the anterior and posterior parts of heart tube: As mentioned in the preceding section, a very regular monophasic potential change was recorded from a localized area in the 7th segment. The same tracing was also found at the anterior end of the 1st segment, where the artery is enlarged in the form of an ampula. The beat frequencies of these two places were three times greater than at other parts of heart. Under the microscope very minute contractions corresponding to the electric changes were found there. A typical electrogram of the 7th segment illustrated in figure 5A consists FIG.5.•@ Action potentials from the 7th segment: A: Repetitive action potential corresponding to the local miniature con- traction. B: A large action potential follow- ing after local contraction of the 7th seg- ment. C-D: Similar tracing led off from the anterior end of heart tube. Many small action potentials coincide with the contrac- tion of anterior end of heart tube, while a larger one coincides with that of the 1st segment. of slow rising phase followed by a rapid spike potential. B, C and D are ex- amples which were all led off from the anterior and posterior ends of the heart tube. One illustrates the small action potential which gradually develops into a large one, while other examples show large number of rapid small action po- tentials of the 7th segment which are preceded by a regular contraction of the 6th segment. 4) Determination of pacemakers by the transection method: In seeking the localization of pacemakers the transection method (1, 8b) was used. The heart 154 H. IRISAWA ET AL. tube was severed into two parts with a sharp razor-blade at the level of the 3rd segment to the4th segment. From table1it is clear that both divided parts beat with nearly equal rhythm. It is suggested that the heart of cicada seems to have two or more than two pacemakers. When the heart tube was severed into more than two parts, the following tendencies were noticed. TABLE1 TABLE2 TABLE3 Each number shows heart beat (frequency) per1minute. E. H.: exposed heart S. S. A.: semisection of alary muscle B. S. A.: bisection of alary muscle the Roman numerals (1st line: segment number, 1st column: case number) √:location sectioning. a) One group of the results is illustrated in table2, where the frequency of the2nd segment is quite similar to that of the normal heart beat. This fact shows that the2nd segment plays a dominant role in pacemaker activity. Es- pecially in the cases II, III and IV, after the heart tube was severed at the level of the3rd segment, the separated posterior part completly ceased to beat, while the anterior part continued to contract with a frequency equal to that of the intact heart. b) Another group (table3) shows a quite different tendency from the former. The examples here show that the posterior end beats more fre- ELECTROGRAMS OF THE CICADA'S HEART 155 quently than the anterior end, after the heart tube was divided into several parts. Among these posterior segments, the7th segment seems to play a domi- nant role in the pacemaker activity. This table also shows that the frequency of the contraction of the posterior segment does not show any remarkable difference from that of the intact heart. From these three tables, it seems that there are two pacemakers in the heart of cicada, one at the2nd segment and another at the7th segment. As the heart was further severed into smaller pieces, they usually ceased beating. The smallest unit required to maintain its rhythmical contraction is illustrated schematically in figure6. When the heart tube was severed like in figure A and B, the beat stopped, while in the case of C, this section can con- tinue to beat, with a frequency of from21to30per minute. This fact indicates that at least one complete segment is needed to keep the rhythmical contraction. FIG.6.•@ Schematic sketch of heart tube, showing the places of transverse section. With cuts as in A and B the middle segment ceases to beat. With cuts as in C the middle segment continues beating (see Text). 5) Reversal of heart beat: In many instances the contraction waves usually progress forwards, that is, from the posterior to anterior end. However, after a while the contraction reverses its direction and begins to progress backwards, It has already been described by Wigglesworth (22) that when the insect is at rest, its blood pressure is nearly equall or even less than the atomospheric pressure.
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