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Sound Signalling in Orthoptera

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Sound Signalling in Orthoptera Open Research Online The Open University’s repository of research publications and other research outputs Sound Signalling in Orthoptera Book Section How to cite: Robinson, David J. and Hall, Marion J. (2002). Sound Signalling in Orthoptera. In: Evans, Peter ed. Advances in Insect Physiology, Volume 29. Elsevier Ltd, pp. 151–278. For guidance on citations see FAQs. c [not recorded] Version: [not recorded] Link(s) to article on publisher’s website: http://dx.doi.org/doi:10.1016/S0065-2806(02)29003-7 Copyright and Moral Rights for the articles on this site are retained by the individual authors and/or other copyright owners. For more information on Open Research Online’s data policy on reuse of materials please consult the policies page. oro.open.ac.uk Sound Signalling in Orthoptera David J Robinson1 and Marion J Hall Department of Biological Sciences, The Open University, Milton Keynes, MK7 6AA, UK Abstract ................................................................................................................................................2 1 Introduction .......................................................................................................................................2 2 Mating systems..................................................................................................................................3 2.1 Variation in mating systems .......................................................................................................3 2.2 Patterns of calling.......................................................................................................................4 3 Songs and signals ..............................................................................................................................5 3.1 Intensity, distance and size .........................................................................................................6 3.2 Mechanisms of sound production...............................................................................................6 3.3 Pattern generation in crickets .....................................................................................................8 3.4 Changes in sound signals with age.............................................................................................8 3.5 Vibratory communication...........................................................................................................9 3.6 Sex differences ...........................................................................................................................9 3.7 Temperature effects....................................................................................................................9 3.8 The energetic costs of calling ...................................................................................................10 4 Hearing and ears..............................................................................................................................10 4.1 Structure of hearing organs.......................................................................................................11 4.2 Auditory receptor organs in the tibia........................................................................................17 4.3 Tonotopic organization of receptor projections........................................................................20 4.4 Directional hearing ...................................................................................................................20 5 Analysis...........................................................................................................................................21 5.1 Song analysis............................................................................................................................22 5.2 Identified auditory interneurons in the nervous system............................................................24 5.3 Symmetry and asymmetry........................................................................................................31 6 Information content of signals.........................................................................................................32 6.1 Components of sound signals...................................................................................................32 6.2 Effects of the environment on sound signals............................................................................33 6.3 Species recognition...................................................................................................................34 6.4 Recognition of sex....................................................................................................................35 6.5 Mate Location...........................................................................................................................36 6.6 Mate choice ..............................................................................................................................36 7 Exploitation of heterospecific sounds..............................................................................................40 7.1 Defences against acoustically-orienting predators in general...................................................41 7.2 Defences against parasitoids.....................................................................................................43 7.3 Defences against bats ...............................................................................................................46 7.4 Sexual versus natural selection.................................................................................................50 7.5 Evolution of predator avoidance mechanisms..........................................................................50 8 Cooperation and competition between males..................................................................................51 8.1 Spacing, aggregating and fighting ............................................................................................51 8.2 Choruses ...................................................................................................................................52 8.3 Satellite males and silent searching..........................................................................................55 9 New directions.................................................................................................................................55 Acknowledgements ............................................................................................................................56 References ..........................................................................................................................................57 1 This review was completed while DJR was a visiting professor at the University of Nagoya, Japan. ADVANCES IN INSECT PHYSIOLOGY VOL. 29, pp. 151-278 (2002) 2 D. J. ROBINSON AND M. J. HALL Abstract The sounds produced by orthopteran insects are very diverse. They are widely studied for the insight they give into acoustic behaviour and the biophysical aspects of sound production and hearing, as well as the transduction of sound to neural signals in the ear and the subsequent processing of information in the central nervous system. The study of sound signalling is a multidisciplinary area of research, with a strong physiological contribution. This review considers recent research in physiology and the links with related areas of acoustic work on the Orthoptera. 1 Introduction Many orthopteran insects are noisy, a lot are large enough for physiological and neurobiological research, and some at least are of economic significance. So, it is not surprising that orthopteran insects are popular with entomologists and much of the research on insect sound has been carried out using members of this order. Their large size relative to many other insects has enabled detailed studies of the nervous system and the ear, and analysis of hearing at the level of identifiable single neurons has been possible. As information has accumulated about the process of detection and analysis of both intraspecific and interspecific signals, from laboratory studies, it has been possible to carry out more detailed analysis of the behaviour of animals under natural conditions and this has included recording from identified interneurons in the field. Developments in the study of behavioural ecology, for example the interest in asymmetry and its role in mate choice, and the evolution of animal signalling, have opened up new areas of research, such as the degree to which the auditory system shows asymmetry at the neuronal level and the physiological costs of signals. The application of cladistic methods to the study of the evolution of the group has thrown up interesting problems, as the similarities between the auditory systems of members of the group at the neuronal level may not imply a common ancestry. As the process of integrating the results from physiological, behavioural and evolutionary studies gathers pace, it seems an opportune time to review recent advances in the study of the physiology of hearing and sound production in the Orthoptera. This is an extensive field of endeavour. In this review, we concentrate on the most recent physiological research and summarize the results and conclusions reported. In addition, we review some recent work in related areas of acoustic behaviour, physiological ecology and the
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