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The Evolution, Ontogeny and Quantitative Control of the Settling Movements of Some New World Saturniid Moths, With Some Comments On Distance Communication By Honey-Bees

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[1. The movement of settling into the rest-position in the New World Hemileucine Saturniine moths is accompanied by rhythmic side-to-side oscillations of the entire animal. The strength of this rocking response can be readily quantified as the number of complete oscillations performed, which may be counted visually. It has been studied comparatively in 52 species of Hemileucine and Citheroniine Saturniids. 2. Homologues of the rocking response are found in the Citheroniinae, which must therefore be more closely related to the Hemileucinae than to the Rhescyntinae, which do not possess them. The rocking responses of the Citheroniinae show greater interspecific diversity, as well as a wider range of variation in their co-ordination within species. The overall evolutionary trend has been towards the reduction and ultimately the elimination of the rocking response. The origin and function of the response remain unknown. 3. A quantitative study has been made of the factors influencing the strength of the rocking response in adult Automeris aurantiaca Weym., allowed to emerge and stored at 20° C. All experimental procedures were conducted at 26-29° C. Some preliminary experiments with Automeris memusae Walker conducted at 26° C are also reported. Flight responses were elicited by suspending the moths in an abdominal clip, and evoking the tarsal reflex; settling responses were studied by placing them on a standardised vertical surface, and observing the strength of the subsequent rocking responses. 4. The strength of the rocking responses given by adult moths once the process of emergence and wing-expansion has been completed is influenced by three factors. When separated, these prove to be : (i) The duration of the preceding period of flight, to which it bears a direct linear relation of the form y = a + bx. In isolation from (ii) and (iii) this effect is termed the long-term relationship. (ii) The proximity of the flight response to the act of settling. When the flight response closely precedes the rocking response in time, the latter is reduced in strength. Further, the build-up of flight excitation over a short period is reflected in a decremental course of the rocking response. These effects are described in terms of a short-term relationship between the factors responsible for mediating the acts of flight and settling. (iii) The age from eclosion, which reduces both a and b in the equation given under (i) above, most steeply near to the act of eclosion. The rocking response is closely linked with the mechanism of eclosion and wing-expansion, and it is argued that part of the changes in the rocking response that occur with age may be the result of changes in central activity relating to the initiation and delimitation in time of these two acts. 5. The long-term interaction between flight and the rocking response is examined in detail. It is shown that (i) acts other than flight intercalated between flight and the rocking response do not affect the strength of the latter. (ii) In the absence of renewed flight, and with an adequate interval between tests to eliminate the short-term interaction, the strength of the rocking response remains stable for at least 90 *minutes after flight. (iii) A variety of procedures fail either to interfere with this stability, or to prevent the incremental course of the rocking response with flight performance; they comprise (a) the perfusion of preparations from which the abdomen has been removed before flight with Ringer's solutions, alone, and with added glucose or trehalose up to concentrations of 60 gms/1.; (b) the removal of the antennae, including Johnston's antennal organ, and Eltringham's organ together with the abdomen before flight; (c) the cutting of the indirect flight muscles and bilateral excision of the wing-base areas before flight. The registration of flight performance and its expression quantitatively in terms of the rocking response must, therefore, be mediated by central nervous interaction. 6. 'Neutral posture', previously interpreted as a state of sustained reciprocal inhibition between the central nervous systems mediating flight and settling is re-examined. A simple statistical test of this assertion is designed and applied, and shown to provide reasonable proof of this interpretation. 7. The ontogeny of adult behaviour in the last 7 days of the pupal development of Automeris memusae is described. The Type II Rhythmic Display develops relatively early in ontogeny. At a stage a few hours before eclosion behaviour is primed, both display and flight behaviour are transiently suppressed. The act of eclosion and the post-eclosion sequence are primed out of phase with the emergence rhythm, and priming is not complete until the rocking response has fully developed. Once this development has taken place, eclosion and wing-expansion may be induced artificially in moths dissected from the pupa by stimulating a brief burst of walking. The development of the rocking response in the 8-12 hours during which the act of eclosion is primed follows a course opposite to that of its rapid decline when the act of eclosion has been completed. 8. Evidence is presented which suggests that the system mediating the strength of the rocking response is sensitive to persistent states of excitation in the central neural mechanisms maintaining and facilitating flight, the modulation of which is considered to be responsible for flight initiation. This working hypothesis is developed and shown to be consistent with the changes in behaviour with age, and with the properties of the short and long-term interactions. 9. Some additional data are given on the relations between oviposition behaviour in virgin females, and the settling and flight responses. Precurrent flight performance in old virgin females releases oviposition, which in turn inhibits settling. The integrity of the abdominal ganglia is necessary for this inhibition, for removal of the abdomen abolishes it. 10. The similarity of the present results to those of recent work on the waggle phase of the distance communication dance of the honey-bee is discussed. It is shown that an hypothesis which regards the relation between the bee's flight performance and the subsequent intensity of the waggle component of the dance as being mediated by central nervous interactions is in better accord with the known facts than the previous theories which have been proposed based on hypothetical exteroceptive or interoceptive feedback systems., 1. The movement of settling into the rest-position in the New World Hemileucine Saturniine moths is accompanied by rhythmic side-to-side oscillations of the entire animal. The strength of this rocking response can be readily quantified as the number of complete oscillations performed, which may be counted visually. It has been studied comparatively in 52 species of Hemileucine and Citheroniine Saturniids. 2. Homologues of the rocking response are found in the Citheroniinae, which must therefore be more closely related to the Hemileucinae than to the Rhescyntinae, which do not possess them. The rocking responses of the Citheroniinae show greater interspecific diversity, as well as a wider range of variation in their co-ordination within species. The overall evolutionary trend has been towards the reduction and ultimately the elimination of the rocking response. The origin and function of the response remain unknown. 3. A quantitative study has been made of the factors influencing the strength of the rocking response in adult Automeris aurantiaca Weym., allowed to emerge and stored at 20° C. All experimental procedures were conducted at 26-29° C. Some preliminary experiments with Automeris memusae Walker conducted at 26° C are also reported. Flight responses were elicited by suspending the moths in an abdominal clip, and evoking the tarsal reflex; settling responses were studied by placing them on a standardised vertical surface, and observing the strength of the subsequent rocking responses. 4. The strength of the rocking responses given by adult moths once the process of emergence and wing-expansion has been completed is influenced by three factors. When separated, these prove to be : (i) The duration of the preceding period of flight, to which it bears a direct linear relation of the form y = a + bx. In isolation from (ii) and (iii) this effect is termed the long-term relationship. (ii) The proximity of the flight response to the act of settling. When the flight response closely precedes the rocking response in time, the latter is reduced in strength. Further, the build-up of flight excitation over a short period is reflected in a decremental course of the rocking response. These effects are described in terms of a short-term relationship between the factors responsible for mediating the acts of flight and settling. (iii) The age from eclosion, which reduces both a and b in the equation given under (i) above, most steeply near to the act of eclosion. The rocking response is closely linked with the mechanism of eclosion and wing-expansion, and it is argued that part of the changes in the rocking response that occur with age may be the result of changes in central activity relating to the initiation and delimitation in time of these two acts. 5. The long-term interaction between flight and the rocking response is examined in detail. It is shown that (i) acts other than flight intercalated between flight and the rocking response do not affect the strength of the latter. (ii) In the absence of renewed flight, and with an adequate interval between tests to eliminate the short-term interaction, the strength of the rocking response remains stable for at least 90 *minutes after flight. (iii) A variety of procedures fail either to interfere with this stability, or to prevent the incremental course of the rocking response with flight performance; they comprise (a) the perfusion of preparations from which the abdomen has been removed before flight with Ringer's solutions, alone, and with added glucose or trehalose up to concentrations of 60 gms/1.; (b) the removal of the antennae, including Johnston's antennal organ, and Eltringham's organ together with the abdomen before flight; (c) the cutting of the indirect flight muscles and bilateral excision of the wing-base areas before flight. The registration of flight performance and its expression quantitatively in terms of the rocking response must, therefore, be mediated by central nervous interaction. 6. 'Neutral posture', previously interpreted as a state of sustained reciprocal inhibition between the central nervous systems mediating flight and settling is re-examined. A simple statistical test of this assertion is designed and applied, and shown to provide reasonable proof of this interpretation. 7. The ontogeny of adult behaviour in the last 7 days of the pupal development of Automeris memusae is described. The Type II Rhythmic Display develops relatively early in ontogeny. At a stage a few hours before eclosion behaviour is primed, both display and flight behaviour are transiently suppressed. The act of eclosion and the post-eclosion sequence are primed out of phase with the emergence rhythm, and priming is not complete until the rocking response has fully developed. Once this development has taken place, eclosion and wing-expansion may be induced artificially in moths dissected from the pupa by stimulating a brief burst of walking. The development of the rocking response in the 8-12 hours during which the act of eclosion is primed follows a course opposite to that of its rapid decline when the act of eclosion has been completed. 8. Evidence is presented which suggests that the system mediating the strength of the rocking response is sensitive to persistent states of excitation in the central neural mechanisms maintaining and facilitating flight, the modulation of which is considered to be responsible for flight initiation. This working hypothesis is developed and shown to be consistent with the changes in behaviour with age, and with the properties of the short and long-term interactions. 9. Some additional data are given on the relations between oviposition behaviour in virgin females, and the settling and flight responses. Precurrent flight performance in old virgin females releases oviposition, which in turn inhibits settling. The integrity of the abdominal ganglia is necessary for this inhibition, for removal of the abdomen abolishes it. 10. The similarity of the present results to those of recent work on the waggle phase of the distance communication dance of the honey-bee is discussed. It is shown that an hypothesis which regards the relation between the bee's flight performance and the subsequent intensity of the waggle component of the dance as being mediated by central nervous interactions is in better accord with the known facts than the previous theories which have been proposed based on hypothetical exteroceptive or interoceptive feedback systems.]

Affiliations: 1: Department of Zoology and Comparative Anatomy, University College London, Canal Zone Biological Area, Drawer C. Balboa, Canal Zone (Panama)

10.1163/156853960X00160
/content/journals/10.1163/156853960x00160
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/content/journals/10.1163/156853960x00160
1960-01-01
2016-12-08

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