Review Article
Mechanics and aerodynamics of insect flight control
- GRAHAM K. TAYLOR
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- Published online by Cambridge University Press:
- 28 November 2001, pp. 449-471
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Insects have evolved sophisticated flight control mechanisms permitting a remarkable range of manoeuvres. Here, I present a qualitative analysis of insect flight control from the perspective of flight mechanics, drawing upon both the neurophysiology and biomechanics literatures. The current literature does not permit a formal, quantitative analysis of flight control, because the aerodynamic force systems that biologists have measured have rarely been complete and the position of the centre of gravity has only been recorded in a few studies. Treating the two best-known insect orders (Diptera and Orthoptera) separately from other insects, I discuss the control mechanisms of different insects in detail. Recent experimental studies suggest that the helicopter model of flight control proposed for Drosophila spp. may be better thought of as a facultative strategy for flight control, rather than the fixed (albeit selected) constraint that it is usually interpreted to be. On the other hand, the so-called ‘constant-lift reaction’ of locusts appears not to be a reflex for maintaining constant lift at varying angles of attack, as is usually assumed, but rather a mechanism to restore the insect to pitch equilibrium following a disturbance. Differences in the kinematic control mechanisms used by the various insect orders are related to differences in the arrangement of the wings, the construction of the flight motor and the unsteady mechanisms of lift production that are used. Since the evolution of insect flight control is likely to have paralleled the evolutionary refinement of these unsteady aerodynamic mechanisms, taxonomic differences in the kinematics of control could provide an assay of the relative importance of different unsteady mechanisms. Although the control kinematics vary widely between orders, the number of degrees of freedom that different insects can control will always be limited by the number of independent control inputs that they use. Control of the moments about all three axes (as used by most conventional aircraft) has only been proven for larger flies and dragonflies, but is likely to be widespread in insects given the number of independent control inputs available to them. Unlike in conventional aircraft, however, insects' control inputs are likely to be highly non-orthogonal, and this will tend to complicate the neural processing required to separate the various motions.
Cephalopod chromatophores: neurobiology and natural history
- J. B. MESSENGER
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- Published online by Cambridge University Press:
- 28 November 2001, pp. 473-528
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The chromatophores of cephalopods differ fundamentally from those of other animals: they are neuromuscular organs rather than cells and are not controlled hormonally. They constitute a unique motor system that operates upon the environment without applying any force to it. Each chromatophore organ comprises an elastic sacculus containing pigment, to which is attached a set of obliquely striated radial muscles, each with its nerves and glia. When excited the muscles contract, expanding the chromatophore; when they relax, energy stored in the elastic sacculus retracts it. The physiology and pharmacology of the chromatophore nerves and muscles of loliginid squids are discussed in detail. Attention is drawn to the multiple innervation of dorsal mantle chromatophores, of crucial importance in pattern generation. The size and density of the chromatophores varies according to habit and lifestyle. Differently coloured chromatophores are distributed precisely with respect to each other, and to reflecting structures beneath them. Some of the rules for establishing this exact arrangement have been elucidated by ontogenetic studies. The chromatophores are not innervated uniformly: specific nerve fibres innervate groups of chromatophores within the fixed, morphological array, producing ‘physiological units’ expressed as visible ‘chromatomotor fields’.
The chromatophores are controlled by a set of lobes in the brain organized hierarchically. At the highest level, the optic lobes, acting largely on visual information, select specific motor programmes (i.e. body patterns); at the lowest level, motoneurons in the chromatophore lobes execute the programmes, their activity or inactivity producing the patterning seen in the skin. In Octopus vulgaris there are over half a million neurons in the chromatophore lobes, and receptors for all the classical neurotransmitters are present, different transmitters being used to activate (or inhibit) the different colour classes of chromatophore motoneurons. A detailed understanding of the way in which the brain controls body patterning still eludes us: the entire system apparently operates without feedback, visual or proprioceptive.
The gross appearance of a cephalopod is termed its body pattern. This comprises a number of components, made up of several units, which in turn contains many elements: the chromatophores themselves and also reflecting cells and skin muscles. Neural control of the chromatophores enables a cephalopod to change its appearance almost instantaneously, a key feature in some escape behaviours and during agonistic signalling. Equally important, it also enables them to generate the discrete patterns so essential for camouflage or for signalling. The primary function of the chromatophores is camouflage. They are used to match the brightness of the background and to produce components that help the animal achieve general resemblance to the substrate or break up the body's outline. Because the chromatophores are neurally controlled an individual can, at any moment, select and exhibit one particular body pattern out of many. Such rapid neural polymorphism (‘polyphenism’) may hinder search-image formation by predators. Another function of the chromatophores is communication. Intraspecific signalling is well documented in several inshore species, and interspecific signalling, using ancient, highly conserved patterns, is also widespread. Neurally controlled chromatophores lend themselves supremely well to communication, allowing rapid, finely graded and bilateral signalling.
Fig-eating by vertebrate frugivores: a global review
- MIKE SHANAHAN, SAMSON SO, STEPHEN G. COMPTON, RICHARD CORLETT
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- Published online by Cambridge University Press:
- 28 November 2001, pp. 529-572
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The consumption of figs (the fruit of Ficus spp.; Moraceae) by vertebrates is reviewed using data from the literature, unpublished accounts and new field data from Borneo and Hong Kong. Records of frugivory from over 75 countries are presented for 260 Ficus species (approximately 30% of described species). Explanations are presented for geographical and taxonomic gaps in the otherwise extensive literature. In addition to a small number of reptiles and fishes, 1274 bird and mammal species in 523 genera and 92 families are known to eat figs. In terms of the number of species and genera of fig-eaters and the number of fig species eaten we identify the avian families interacting most with Ficus to be Columbidae, Psittacidae, Pycnonotidae, Bucerotidae, Sturnidae and Lybiidae. Among mammals, the major fig-eating families are Pteropodidae, Cercopithecidae, Sciuridae, Phyllostomidae and Cebidae. We assess the role these and other frugivores play in Ficus seed dispersal and identify fig-specialists. In most, but not all, cases fig specialists provide effective seed dispersal services to the Ficus species on which they feed. The diversity of fig-eaters is explained with respect to fig design and nutrient content, phenology of fig ripening and the diversity of fig presentation. Whilst at a gross level there exists considerable overlap between birds, arboreal mammals and fruit bats with regard to the fig species they consume, closer analysis, based on evidence from across the tropics, suggests that discrete guilds of Ficus species differentially attract subsets of sympatric frugivore communities. This dispersal guild structure is determined by interspecific differences in fig design and presentation. Throughout our examination of the fig-frugivore interaction we consider phylogenetic factors and make comparisons between large-scale biogeographical regions. Our dataset supports previous claims that Ficus is the most important plant genus for tropical frugivores. We explore the concept of figs as keystone resources and suggest criteria for future investigations of their dietary importance. Finally, fully referenced lists of frugivores recorded at each Ficus species and of Ficus species in the diet of each frugivore are presented as online appendices. In situations where ecological information is incomplete or its retrieval is impractical, this valuable resource will assist conservationists in evaluating the role of figs or their frugivores in tropical forest sites.