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Hawks, doves, and mixed-symmetry games.

The hawk-dove game has proved to be an important tool for understanding the role of aggression in social interactions. Here, the game is presented in a more general form (GHD) to facilitate analyses of interactions between individuals that may differ in "size", where size is interpreted as a surrogate for resource holding power. Three different situations are considered, based on the availability and use of information that interacting individuals have about their sizes: the classical symmetric case, in which no information about sizes is used, the asymmetric case, in which the individuals know their relative sizes and thus their chances of prevailing in combat, and a mixed-symmetry case, in which each individual only knows its own size (or only knows its opponent's size). I describe and use some recently developed methods for multitype games-evolutionary games involving two or more categories of players. With these methods and others, the evolutionarily stable strategies (ESSs) that emerge for the three different cases are identified and compared. A proof of the form and uniqueness of the ESS for the mixed-symmetry case is presented. In this situation, one size category at most can play a mixed strategy; larger individuals are aggressive and smaller individuals are not. As the number of size categories approaches infinity and the size distribution becomes continuous, there is a threshold size, above which all individuals are aggressive, and below which they are not.

Aggression↗

Vision complements echolocation in an aerial-hawking bat.

The northern bat Eptesicus nilssonii normally hunts flying insects in the air using frequency-modulated echolocation calls. It is also known to detect and catch visually conspicuous prey (white moths) hovering low among grass stalks. To overcome the problem with acoustic clutter from the grass, which interferes with target echo detection, the bats make use of visual cues in addition to those of echolocation. We therefore investigated the minimum size of prey that the bats could distinguish by using vision, by presenting the bats with different sized dead and spread moths. We found that vision increased the chance of detection only when the moths had a wingspan of at least 5 cm. Smaller targets were detected using echolocation alone. The mean detection range was 3.5 m, suggesting that the bats need a visual acuity of 49' of arc to detect the prey. This is consistent with results of optomotor response tests and counts of retinal ganglion cells in closely related species. Our results suggest that the visual acuity of Eptesicus bats may not be adequate for prey detection under normal conditions, but that the bats can use vision when the prey is unusually large and conspicuous. The northern bats display a flexibility in prey detection techniques not previously recognised among aerial-hawking bats and they are able to use their full visual capacity in the field.

Animals↗

Nectar feeding by the hovering hawk moth Macroglossum stellatarum: intake rate as a function of viscosity and concentration of sucrose solutions.

Although nectar feeding in insects has long been studied, the knowledge of the effect of nectar energy content on the ingestion dynamics separately from the viscosity of the fluid is very limited. To determine the effects of both factors on the feeding behavior of the hovering hawk moth Macroglossum stellatarum, we developed a method to independently manipulate sucrose concentrations and viscosity. The intake rate was analyzed as a function of sucrose concentration, the concentration at constant viscosity (kept constant by adding tylose, an inert polysaccharide), and of the different viscosities of a 30% weight/weight (w/w) sucrose solution (by adding different amounts of tylose). By increasing the concentration, and thus its viscosity, the solution intake rate (in microl s (-1)) decreased beyond a 20% w/w sucrose solution. For a 30% sucrose solution, the intake rate decreased with increasing viscosity. At constant viscosity, the solution intake rate decreased beyond a 30% w/w sucrose solution. However, if we considered the quantity of sucrose ingested per unit time (sucrose intake rate), the same fitted maximum was attained for both series in which the sucrose concentration changed (33.6% w/w). Results suggest that the gustatory input affects the dynamics of fluid ingestion separately from the viscosity.

Animals↗

Neural correlates of habituation of the proleg withdrawal reflex in larvae of the hawk moth, Manduca sexta.

The larval proleg withdrawal reflex of the hawk moth, Manduca sexta, exhibits robust habituation. This reflex is evoked by deflecting one or more mechanosensory planta hairs on a proleg tip. We examined neural correlates of habituation in an isolated proleg preparation consisting of one proleg and its segmental ganglion. Repeated deflection of a single planta hair caused a significant decrease in the number of action potentials evoked in the proleg motor nerve (which carries the axons of proleg retractor motor neurons). Significant response decrement was seen for interstimulus intervals of 10 s, 60 s and 5 min. Response decrement failed to occur in the absence of repetitive stimulation, the decremented response recovered spontaneously following a rest, and electrical stimulation of a body wall nerve facilitated the decremented response (a neural correlate of dishabituation). Adaptation of sensory neuron responses occurred during repeated hair deflections. However, when adaptation was eliminated by direct electrical stimulation of sensory neurons, the response in the proleg motor nerve still decreased significantly. Muscle recordings indicated that the response of an identified proleg retractor motor neuron decreased significantly during habituation training. Thus, habituation of the proleg withdrawal reflex includes a central component that is apparent at the level of a single motor neuron.

Animals↗

Visual position stabilization in the hummingbird hawk moth, Macroglossum stellatarum L. II. Electrophysiological analysis of neurons sensitive to wide-field image motion.

Response properties of neurons in the cervical connectives of the hummingbird hawk moth, Macroglossum stellatarum L., were determined. All neurons described in this account respond directionally selectively to motion in large parts of the visual field of either eye. They respond maximally to bilateral stimulation, preferring either motion as induced on the eyes during translatory movements of the animal or when it turns around one of its body axes. Cells most sensitive to rotational motion either respond best to rotation of the patterns around the vertical axis of the animal or around its longitudinal body axis. Neurons most sensitive to translational pattern motion respond best to either simulated translations of the animal along its vertical or along an oblique axis. Most types of neurons respond tonically and do not habituate. The sensitivity to motion stimuli is not evenly distributed within the receptive field of any investigated neuron. Part of these neurons might play a role in visual position and course stabilization.

Animals↗

Expression of UV-, blue-, long-wavelength-sensitive opsins and melatonin in extraretinal photoreceptors of the optic lobes of hawk moths.

Lepidopterans display biological rhythms associated with egg laying, eclosion and flight activity but the photoreceptors that mediate these behavioural patterns are largely unknown. To further our progress in identifying candidate light-input channels for the lepidopteran circadian system, we have developed polyclonal antibodies against ultraviolet (UV)-, blue- and extraretinal long-wavelength (LW)-sensitive opsins and examined opsin immunoreactivity in the adult optic lobes of four hawk moths, Manduca sexta, Acherontia atropos, Agrius convolvuli and Hippotion celerio. Outside the retina, UV and blue opsin protein expression is restricted to the adult stemmata, with no apparent expression elsewhere in the brain. Melatonin, which is known to have a seasonal influence on reproduction and behaviour, is expressed with opsins in adult stemmata together with visual arrestin and chaoptin. By contrast, the LW opsin protein is not expressed in the retina or stemmata but rather exhibits a distinct and widespread distribution in dorsal and ventral neurons of the optic lobes. The lamina, medulla, lobula and lobula plate, accessory medulla and adjacent neurons innervating this structure also exhibit strong LW opsin immunoreactivity. Together with the adult stemmata, these neurons appear to be functional photoreceptors, as visual arrestin, chaoptin and melatonin are also co-expressed with LW opsin. These findings are the first to suggest a role for three spectrally distinct classes of opsin in the extraretinal detection of changes in ambient light and to show melatonin-mediated neuroendocrine output in the entrainment of sphingid moth circadian and/or photoperiodic rhythms.

Animals↗

Development of adult thoracic leg muscles during metamorphosis of the hawk moth Manduca sexta.

During metamorphosis, the larval thoracic legs of the hawk moth Manduca sexta are replaced by a new set of adult legs. The larval leg motoneurons persist to innervate new adult muscles, and the motor terminals remain within the developing adult legs. Here we describe the fate of the larval leg muscles and the origin of new muscles within the adult legs. During the larval instars, large and small nuclei proliferate within leg muscle fibers. Near the end of the larval stage a subset of the small nuclei undergo a wave of proliferation, as indicated by the incorporation of 5-bromodeoxyuridine, whereas other nuclei die. However, none of the larval leg muscles fibers persist to serve as templates for adult muscle formation, and there was no evidence for persistence of larval myonuclei. Migrating myoblasts that are born within aggregate to form adult muscle anlagen at specific production sites within the developing imaginal legs. Intense nuclear proliferation occurs within the anlagen during the early pupal stage, followed by muscle fiber formation and striation. We conclude that adult leg muscles form mainly, if not exclusively, from migrating myoblasts that without the involvement of larval elements.

Animals↗

The insect brain (Na+ + K+)-ATPase. Binding of ouabain in the hawk moth, Manduca sexta.

(1) A quantitative study has been made of the binding of ouabain to the (Na+ + K+)-ATPase in homogenates prepared from brain tissue of the hawk moth, Manduca sexta. The results have been compared to those obtained in bovine brain microsomes. (2) The insect brain (Na+ + K+)-ATPase will bind ouabain either in the presence of Mg2+ and Pi, ('Mg2+, Pi' conditions) or in the presence of Na+, Mg2+, and an adenine nucleotide ('nucleotide' conditions) as is the case for the bovine brain (Na+ + K+)-ATPase. The binding conditions did not alter the total number of receptor sites measured at high ouabain concentrations in either tissue. (3) Potassium ion decreases the affinity (increases the KD) of ouabain to the M. sexta brain (Na+ + K+)-ATPase under both binding conditions. However, ouabain binding is more sensitive to K+ inhibition under the nucleotide conditions. In bovine brain ouabain binding is equally sensitive to K+ inhibition under both conditions. (4) The enzyme-ouabain complex has a rate of dissociation that is 10-fold faster in the M. sexta preparation than in the bovine brain preparation. Because of this, the M. sexta (Na+ + K+)-ATPase has a higher KD for ouabain binding and is less sensitive to inhibition by ouabain than the bovine brain enzyme. (5) This data supports the hypothesis that two different conformational states of the M. sexta (Na+ + K+)-ATPase can bind ouabain.

Animals↗

Bifurcation analysis of a predator-prey model with predators using hawk and dove tactics.

Most classical prey-predator models do not take into account the behavioural structure of the population. Usually, the predator and the prey populations are assumed to be homogeneous, i.e. all individuals behave in the same way. In this work, we shall take into account different tactics that predators can use for exploiting a common self-reproducing resource, the prey population. Predators fight together in order to keep or to have access to captured prey individuals. Individual predators can use two behavioural tactics when they encounter to dispute a prey, the classical hawk and dove tactics. We assume two different time scales. The fast time scale corresponds to the inter-specific searching and handling for the prey by the predators and the intra-specific fighting between the predators. The slow time scale corresponds to the (logistic) growth of the prey population and mortality of the predator. We take advantage of the two time scales to reduce the dimension of the model and to obtain an aggregated model that describes the dynamics of the total predator and prey densities at the slow time scale. We present the bifurcation analysis of the model and the effects of the different predator tactics on persistence and stability of the prey-predator community are discussed.

Animals↗

Can transitive inference evolve in animals playing the hawk-dove game?

What should an individual do if there are no reliable cues to the strength of a competitor when fighting with it for resources? We herein examine the evolutionarily stable strategy (ESS) in the hawk-dove game, if the opponent's resource-holding potential (RHP) can only indirectly be inferred from the outcome of past interactions in the population. The strategies we examined include the classical mixed strategy in which no information on past games is utilized, the 'imprinting' strategy in which a player increases/decreases its aggressiveness if it wins/loses a game, the 'immediate inference' strategy in which a player can infer the strength of those opponents it fought before, and the 'transitive inference' strategy in which a player can infer the strength of a new opponent through a third party with which both players have fought before. Invasibility analysis for each pair of strategies revealed that (i) the transitive-inference strategy can always invade the mixed strategy and the imprinting strategy, and itself refuses invasion by these strategies; (ii) the largest advantage for transitive inference is achieved when the number of games played per individual in one generation is small and when the cost of losing an escalated game is large; (iii) the immediate inference, rather than the transitive inference, can be an ESS if the cost of fighting is small; (iv) a strong linear ranking is established in the population of transitive-inference strategists, though it does not perfectly correlate to the ranking by actual RHPs. We found that the advantage of the transitive inference is not in its ability to correct a misassessment (it is actually the worst in doing so), but in the ability of quickly lining up either incorrect or correct assessments to form a linear dominance hierarchy.

Animals↗

Cognitive requirements for hawk-dove games: A functional analysis for evolutionary design.

Like other social animals, humans play adaptively important games, and current evolutionary theory predicts special-purpose, domain-specific cognitive mechanisms for playing such games. We offer a functional analysis of the information requirements for successfully playing one important social game, the "hawk-dove" conflict-of-interest game, developing new graphic conventions for doing so. In particular, we address the orders of recognition necessary for successfully playing such games, showing that there are adaptive advantages of capacities for first, second, third, and fourth such orders, but no more. We suggest that first-order recognition is not only the most basic in analytic terms but is likely to have been the first to evolve, with subsequent orders added later in evolution.

Journal Article↗

The cost of misinformation in deadly conflicts: Hawk-Dove games and suicidal terrorism.

The hijacking and purposeful crashing of airplanes into the World Trade Center and the Pentagon on September 11, 2001, prompts questions about why the passengers and crew of those airplanes did not act to prevent these attacks, as did at least some passengers on a hijacked flight that crashed in Pennsylvania. We argue, first, that humans have an evolved cognitive bias that leads to the expectation that antagonists hope to survive conflict and, second, that highly credible information to the contrary is needed to overcome this bias. Absent such information, the passengers on at least two airplanes incorrectly interpreted the game being played as a hawk-dove version of a conflict-of-interest game, when it was actually a "suicidal terrorism" variant of that family. Given that other terrorists may have been in the air and ready to act, the airlines' policy of not informing passengers about such events could have risked disabling them from reacting forcefully when force alone was advisable.

Journal Article↗

Oscillations of frequency in Batesian mimics, hawks and doves, and other simple frequency dependent polymorphisms.

It is customary to infer the properties of the internal equilibria produced by frequency dependent selection from the properties of the boundary equilibria (often called the "invasion" criterion). This paper demonstrates that there are some circumstances that there is a truly stable, unique, internal equilibrium. For two alleles with complete dominance, if phenotype fitness declines monotonically with increasing frequency, then the internal point of equal phenotype fitness is the unique internal equilibrium of the genes, and is not unstable; this criterion may also be met if the fitness of one phenotype increases with frequency. It must be truly stable, in the sense of not producing oscillations, if the decline of fitness is linear or convex upwards and no phenotype is lethal at any frequency; the hawk-dove game complies with both conditions, and at least the second condition is likely to be met in most of the models encountered in sociobiology. However, an equilibrium which induces damped oscillations, or perhaps even complex limit cycles, is possible if at least one phenotype can be lethal at high frequency, or if the decline in fitness is strongly curvilinear and concave upwards. One case of curvilinear frequency dependence, a dimorphic batesian mimic with a non-mimetic form, is examined in detail. Although oscillations about the recessive or Y-linked, this will only occur when selection coefficients are very large, and (except for Y-linkage) only if both sexes can be mimetic. As selection is density as well as frequency dependent, such conditions may be produced in the real world by large fluctuations in population size.

Adaptation, Biological↗

The rate of substrate cycling between glucose and glucose 6-phosphate in muscle and fat-body of the hawk moth (Acherontia atropos) at rest and during flight.

1. The rate of substrate cycling between glucose and glucose 6-phosphate was measured in tissues of the hawk moth (Acherontia atropos). 2. The insect was injected with [2-3H,2-14C]glucose, and after periods of time at rest or flying the animal was freeze-clamped. Separation of glucose and hexose monophosphate from the tissues was performed by paper chromatography and t.l.c., and the 3H and 14C radioactivities in these compounds were measured. 3. On the basis of the 3H/14C ratios in these compounds and the measured rate of glycolysis, the rate of cycling was calculated. The rates of cycling were 0.03, 0.10, 0.06 and 3.9 mumol/min per g for fat-body at rest and during flight and for flight muscle at rest and during flight respectively. 4. The marked increase in the cycling rate between glucose and glucose 6-phosphate upon flight contrasts with the finding of Clark, Bloxham, Holland & Lardy [(1973) Biochem. J. 134, 589-597] in the bumble-bee, in which this condition inhibited cycling. It is suggested that the increased rate of cycling increases the sensitivity of glucose phosphorylation to changes in the concentrations of effectors of hexokinase should it be necessary to increase the rate of glycolysis in muscle, for example, to increase power output of the flight muscle for increased speed of flight.

Adipose Tissue↗

A survey on the incidence and cause of ovine abortion in Hawkes Bay.

The results of a postal survey which yielded a 57% response and which examined the incidence and cause of abortions in central Hawkes Bay flocks, is presented. Data were provided from a total of 3,952,000 ewes from 1975-80. Overall, highly significant differences in the incidence of abortion were recorded between properties (range 0-16%); years (1975-0.86%, 1976-0.75%, 1977-0.86%, 1978-2.42%, 1979-2.78%, 1980-2.04%); ewe age (maiden 2-year -5.3%, mixed age -1.6%);flock size (higher incidence in smaller flocks); and winter management systems (of the flocks which were experiencing greater than a 2% abortion rate, 73.2% were being rotationally mob stocked during the winter compared with only 23.8% being set stocked). Bacteriological examination of aborted lambs during 1979 revealed that Campylobacter spp. were involved exclusively. In the 1980 survey, only half of the lambs examined showed Campylobacter infection, the other half yielding Toxoplasma either as pure or mixed infections. The results of a trial which utilised 10,592 2-year ewes demonstrated significant differences between animals vaccinated with a killed Campylobacter vaccine and non-vaccinated controls in the following reproductive parameters: ewes aborting, dry dry ewes, total lambs born and lambs marked.

Journal Article↗

The scavenging behaviour of ferrets (Mustela furo), feral cats (Felis domesticus), possums (Trichosurus vulpecula), hedgehogs (Erinaceus europaeus) and harrier hawks (Circus approximans) on pastoral farmland in New Zealand: implications for bovine tuberculosis transmission.

AIMS: To identify species that scavenge carcasses in pastoral habitats in New Zealand; to determine whether there were interspecific or intraspecific differences in scavenging behaviour and; to document any interspecific or intraspecific interactions occurring at carcasses. METHODS: Scavenging by ferrets (Mustela furo), feral cats (Felis domesticus), possums (Trichosurus vulpecula), hedgehogs (Erinaceus europaeus) and harrier hawks (Circus approximans) was studied from autumn to midwinter on pastoral farmland near Palmerston (45S, 170E), Otago, New Zealand. Time-lapse video recorders and camera lens mounted with infra-red light illumination were used to monitor carcasses of 10 ferrets, 12 possums, 2 hedgehogs and 7 rabbits (Oryctolagus cuniculus) until they were totally scavenged. RESULTS: Ferrets scavenged 5/8 ferret carcasses, 8/9 possum carcasses and 6/7 rabbit carcasses encountered. Feral cats scavenged 3/8 ferret carcasses, 5/7 rabbit carcasses, and 3/8 possum carcasses encountered. Possums scavenged 1/2 ferret carcasses and 3/4 rabbit carcasses encountered. The proportion of encounters resulting in feeding on ferret carcasses differed between ferrets (45.7%) and possums (6.3%), and between possums and cats (29.7%). Similarly, for possum carcasses, differences were found between ferrets (76.6%) and possums (0%), ferrets and cats (60.6%) and possums and cats. No interspecific differences were found in the proportion of encounters that resulted in feeding on rabbit carcasses between ferrets (85.2%), possums (75%) and cats (73.1%). In 8/12 instances of ferrets coming into contact with other ferrets whilst feeding, ferrets fed together at the carcass. On 1 occasion, 4 ferrets were recorded feeding together. In 7/8 instances where cats and ferrets came into contact over carrion, ferrets maintained possession or displaced the cat from the carcass. CONCLUSIONS: Communal carrion feeding by ferrets may facilitate intraspecific and interspecific transmission of bovine tuberculosis (caused by Mycobacterium bovis) by the consumption of contaminated carrion, fighting, or close-contact activities. Cannibalism may be one mechanism by which tuberculosis is transmitted within ferret populations. Our results also suggest that possums may acquire infection from carrion, despite being mainly herbivorous.

Journal Article↗

Nutritional secondary hyperparathyroidism and osteodystrophia fibrosa in a Hodgson's hawk-eagle (Spizaetus nipalensis).

A Hodgson's hawk-eagle (Spizaetus nipalensis) reared by a falconer showed severe weakness with multiple fractures of bone. It had a history of being fed an all-meat diet. Serological examination revealed a hypocalcaemia (72.0 microg/ml), and hypophosphataemia (29.0 microg/ml). Gross and microscopic examinations demonstrated severe osteodystrophia fibrosa (fibrous osteodystrophy) characterized by osteoclastic bone resorption and intertrabecular fibrosis with unmineralized trabecular bone containing a large amount of unmineralized osteoid. There was also hypertrophy and hyperplasia of the parathyroid glands, which is consistent with nutritional secondary hyperparathyroidism.

Animal Feed↗

Hawks and Doves on small-world networks.

We explore the Hawk-Dove game on networks with topologies ranging from regular lattices to random graphs with small-world networks in between. This is done by means of computer simulations using several update rules for the population evolutionary dynamics. We find the overall result that cooperation is sometimes inhibited and sometimes enhanced in those network structures, with respect to the mixing population case. The differences are due to different update rules and depend on the gain-to-cost ratio. We analyze and qualitatively explain this behavior by using local topological arguments.

Journal Article↗