Motivation matters: cheliped extension displays in the hermit crab, Pagurus bernhardus, are honest signals of hunger

Whether animal signals convey honest information is a central evolutionary question, since selection pressures could, in some circumstances, favour dishonesty. A prior study of signalling in hermit crabs proposed that the cheliped extension display of Pagurus bernhardus might represent such an instance of dishonesty. A limitation of this conclusion, however, was that honesty was defined in the context of size assessment, neglecting the potential information that displays might transmit about signallers' variable internal states. Recent analyses of signalling in this same species have shown that its displays provide reliable information about the amount of risk crabs are prepared to tolerate, which therefore might enable signallers to use these displays to honestly convey their motivation to take such risks. Here we test this 'honest advertisement of motivation' hypothesis by varying crabs' need for food and analysing their signalling during simulated feeding conflicts against a model. When crabs were starved for 1-5 days, they dropped significantly in weight. Despite this decrement in resource-holding potential and energy reserves, crabs were more likely to perform cheliped extension displays the longer they were food deprived. Longer-starved crabs, whose subjective resource value was greater, also displayed at a higher rate and were more likely to risk seizing the food from the model. We conclude that cheliped extension is a reliable indicator of crabs' internal state and suggest how this honest signal might operate in conflicts over a variety of other resources in addition to food. We propose that future studies detecting apparent dishonesty should analyse many possible signal-state correlations before concluding a signal is actually dishonest. (c) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Nestling testosterone is associated with begging behaviour and fledging success in the pied flycatcher, Ficedula hypoleuca

Animal signals are hypothesized to be costly in order to honestly reflect individual quality. Offspring solicitation signals given by nestling birds are thought to have evolved to advertise either need or individual quality. We tested the potential role of testosterone (T) in controlling the intensity of these signals by measuring begging behaviour as: (i) duration of the begging display and (ii) maximum height of the begging stretch, and by sampling endogenous T levels in nestling blood. We tested nestling pied flycatchers (Ficedula hypoleuca) using well-established experimental paradigm involving transient food deprivation to encourage begging behaviour and then blood-sampled nestlings at the end of these tests for T levels. Our results show that individual nestlings with the most intense begging displays had the highest circulating levels of T immediately after testing. In addition, we found substantial differences between broods in terms of circulating T. Finally, we found evidence that broods with higher levels of T showed increased fledging success, indicating a benefit for increased T production in nestlings. The potential trade-offs involved in T-mediated begging behaviour are discussed.

Comparing entire colour patterns as birds see them

Colour patterns and their visual backgrounds consist of a mosaic of patches that vary in colour, brightness, size, shape and position. Most studies of crypsis, aposematism, sexual selection, or other forms of signalling concentrate on one or two patch classes (colours), either ignoring the rest of the colour pattern, or analysing the patches separately. We summarize methods of comparing colour patterns making use of known properties of bird eyes. The methods are easily modifiable for other animal visual systems. We present a new statistical method to compare entire colour patterns rather than comparing multiple pairs of patches. Unlike previous methods, the new method detects differences in the relationships among the colours, not just differences in colours. We present tests of the method's ability to detect a variety of kinds of differences between natural colour patterns and provide suggestions for analysis.

Measurement error causes scale-dependent threshold erosion of biological signals in animal movement data

Recent advances in telemetry technology have created a wealth of tracking data available for many animal species moving over spatial scales from tens of meters to tens of thousands of kilometers. Increasingly, such data sets are being used for quantitative movement analyses aimed at extracting fundamental biological signals such as optimal searching behavior and scale-dependent foraging decisions. We show here that the location error inherent in various tracking technologies reduces the ability to detect patterns of behavior within movements. Our analyses endeavored to set out a series of initial ground rules for ecologists to help ensure that sampling noise is not misinterpreted as a real biological signal. We simulated animal movement tracks using specialized random walks known as Lévy flights at three spatial scales of investigation: 100-km, 10-km, and 1-km maximum daily step lengths. The locations generated in the simulations were then blurred using known error distributions associated with commonly applied tracking methods: the Global Positioning System (GPS), Argos polar-orbiting satellites, and light-level geolocation. Deviations from the idealized Lévy flight pattern were assessed for each track after incrementing levels of location error were applied at each spatial scale, with additional assessments of the effect of error on scale-dependent movement patterns measured using fractal mean dimension and first-passage time (FPT) analyses. The accuracy of parameter estimation (Lévy μ, fractal mean D, and variance in FPT) declined precipitously at threshold errors relative to each spatial scale. At 100-km maximum daily step lengths, error standard deviations of ≥10 km seriously eroded the biological patterns evident in the simulated tracks, with analogous thresholds at the 10-km and 1-km scales (error SD ≥ 1.3 km and 0.07 km, respectively). Temporal subsampling of the simulated tracks maintained some elements of the biological signals depending on error level and spatial scale. Failure to account for large errors relative to the scale of movement can produce substantial biases in the interpretation of movement patterns. This study provides researchers with a framework for understanding the limitations of their data and identifies how temporal subsampling can help to reduce the influence of spatial error on their conclusions.

Reciprocal secretion of proteins by the bacterial type III machines of plant and animal pathogens suggests universal recognition of mRNA targeting signals

Bacterial pathogens of both animals and plants use type III secretion machines to inject virulence proteins into host cells. Although many components of the secretion machinery are conserved among different bacterial species, the substrates for their type III pathways are not. The Yersinia type III machinery recognizes some secretion substrates via a signal that is encoded within the first 15 codons of yop mRNA. These signals can be altered by frameshift mutations without affecting secretion of the encoded polypeptides, suggesting a mechanism whereby translation of yop mRNA is coupled to the translocation of newly synthesized polypeptide. We report that the type III machinery of Erwinia chrysanthemi cloned in Escherichia coli recognizes the secretion signals of yopE and yopQ. Pseudomonas syringae AvrB and AvrPto, two proteins exported by the recombinant Erwinia machine, can also be secreted by the Yersinia type III pathway. Mapping AvrPto sequences sufficient for the secretion of reporter fusions in Yersinia revealed the presence of an mRNA secretion signal. We propose that 11 conserved components of type III secretion machines may recognize signals that couple mRNA translation to polypeptide secretion.

Cone signals and activity in myopia and emmetropia

Myopia (short-sightedness) is a common ocular disorder of children and young adults. Studies primarily using animal models have shown that the retina controls eye growth and the outer retina is likely to have a key role. One theory is that the proportion of L (long-wavelength-sensitive) and M (medium-wavelength-sensitive) cones is related to myopia development; with a high L/M cone ratio predisposing individuals to myopia. However, not all dichromats (persons with red-green colour vision deficiency) with extreme L/M cone ratios have high refractive errors. We predict that the L/M cone ratio will vary in individuals with normal trichromatic colour vision but not show a systematic difference simply due to refractive error. The aim of this study was to determine if L/M cone ratios in the central 30° are different between myopic and emmetropic young, colour normal adults. Information about L/M cone ratios was determined using the multifocal visual evoked potential (mfVEP). The mfVEP can be used to measure the response of visual cortex to different visual stimuli. The visual stimuli were generated and measurements performed using the Visual Evoked Response Imaging System (VERIS 5.1). The mfVEP was measured when the L and M cone systems were separately stimulated using the method of silent substitution. The method of silent substitution alters the output of three primary lights, each with physically different spectral distributions to control the excitation of one or more photoreceptor classes without changing the excitation of the unmodulated photoreceptor classes. The stimulus was a dartboard array subtending 30° horizontally and 30° vertically on a calibrated LCD screen. The m-sequence of the stimulus was 215-1. The N1-P1 amplitude ratio of the mfVEP was used to estimate the L/M cone ratio. Data were collected for 30 young adults (22 to 33 years of age), consisting of 10 emmetropes (+0.3±0.4 D) and 20 myopes (–3.4±1.7 D). The stimulus and analysis techniques were confirmed using responses of two dichromats. For the entire participant group, the estimated central L/M cone ratios ranged from 0.56 to 1.80 in the central 3°-13° diameter ring and from 0.94 to 1.91 in the more peripheral 13°-30° diameter ring. Within 3°-13°, the mean L/M cone ratio of the emmetropic group was 1.20±0.33 and the mean was similar, 1.20±0.26, for the myopic group. For the 13°-30° ring, the mean L/M cone ratio of the emmetropic group was 1.48±0.27 and it was slightly lower in the myopic group, 1.30±0.27. Independent-samples t-test indicated no significant difference between the L/M cone ratios of the emmetropic and myopic group for either the central 3°-13° ring (p=0.986) or the more peripheral 13°-30° ring (p=0.108). The similar distributions of estimated L/M cone ratios in the sample of emmetropes and myopes indicates that there is likely to be no association between the L/M cone ratio and refractive error in humans.

Origin of the double- and multi-pulse structure of echolocation signals in Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientialis)

The signals of dolphins and porpoises often exhibit a multi-pulse structure. Here, echolocation signal recordings were made from four geometrically distinct positions of seven Yangtze finless porpoises temporarily housed in a relatively small, enclosed area. Some clicks demonstrated double-pulse, and others multi-pulse, structure. The interpulse intervals between the first and second pulse of the double- and multi-pulse clicks were significantly different among data from the four different positions (p < 0.01, one-way ANOVA). These results indicate that the interpulse interval and structure of the double- and multi-pulse echolocation signals depend on the hydrophone geometry of the animal, and that the double- and multi-pulse structure of echolocation signals in Yangtze finless porpoise is not caused by the phonating porpoise itself, but by the multipath propagation of the signal. Time delays in the 180 degrees phase-shifted surface reflection pulse and the nonphase-shifted bottom reflection pulse of the multi-pulse structures, relative to the direct signal, can be used to calculate the distance to a phonating animal. (c) 2005 Acoustical Society of America.

A method for individual identification of echolocation signals in free-ranging finless porpoises carrying data loggers

Echolocation click events of a free-ranging juvenile and an adult finless porpoise (Neophocaena phocaenoides) were recorded with an acoustic data logger. Additionally, dive depth and swim speed of the juvenile were recorded with a behavior data logger. Echoes of echolocation signals from the water surface were clearly detected in shallow dives approximately less than 2 m. The delay time between a surface echo and a direct signal corresponded with the two-way transmission time for the animal's depth, indicating that the signals originated from the animal wearing the data loggers. The finless porpoises produced echolocation signals frequently and were thought to be able to detect their depth by listening to echoes from the water surface. (C) 2000 Acoustical Society of America. [S0001-4966(00)01609-X].

Micro Insert: A Prototype Full-Ring PET Device for Improving the Image Resolution of a Small-Animal PET Scanner

A full-ring PET insert device should be able to enhance the image resolution of existing small-animal PET scanners. Methods: The device consists of 18 high-resolution PET detectors in a cylindric enclosure. Each detector contains a cerium-doped lutetium oxyorthosilicate array (12 x 12 crystals, 0.72 x 1.51 x 3.75 mm each) coupled to a position-sensitive photomultiplier tube via an optical fiber bundle made of 8 x 16 square multiclad fibers. Signals from the insert detectors are connected to the scanner through the electronics of the disabled first ring of detectors, which permits coincidence detection between the 2 systems. Energy resolution of a detector was measured using a Ge-68 point source, and a calibrated 68Ge point source stepped across the axial field of view (FOV) provided the sensitivity profile of the system. A Na-22 point source imaged at different offsets from the center characterized the in-plane resolution of the insert system. Imaging was then performed with a Derenzo phantom filled with 19.5 MBq of F-18-fluoride and imaged for 2 h; a 24.3-g mouse injected with 129.5 MBq of F-18-fluoride and imaged in 5 bed positions at 3.5 h after injection; and a 22.8-g mouse injected with 14.3 MBq of F-18-FDG and imaged for 2 h with electrocardiogram gating. Results: The energy resolution of a typical detector module at 511 keV is 19.0% +/- 3.1 %. The peak sensitivity of the system is approximately 2.67%. The image resolution of the system ranges from 1.0- to 1.8-mm full width at half maximum near the center of the FOV, depending on the type of coincidence events used for image reconstruction. Derenzo phantom and mouse bone images showed significant improvement in transaxial image resolution using the insert device. Mouse heart images demonstrated the gated imaging capability of the device. Conclusion: We have built a prototype full-ring insert device for a small-animal PET scanner to provide higher-resolution PET images within a reduced imaging FOV. Development of additional correction techniques are needed to achieve quantitative imaging with such an insert.

Mix it and fix it: functions of composite olfactory signals in ring-tailed lemurs.

Animals communicating via scent often deposit composite signals that incorporate odorants from multiple sources; however, the function of mixing chemical signals remains understudied. We tested both a 'multiple-messages' and a 'fixative' hypothesis of composite olfactory signalling, which, respectively, posit that mixing scents functions to increase information content or prolong signal longevity. Our subjects-adult, male ring-tailed lemurs (Lemur catta)-have a complex scent-marking repertoire, involving volatile antebrachial (A) secretions, deposited pure or after being mixed with a squalene-rich paste exuded from brachial (B) glands. Using behavioural bioassays, we examined recipient responses to odorants collected from conspecific strangers. We concurrently presented pure A, pure B and mixed A + B secretions, in fresh or decayed conditions. Lemurs preferentially responded to mixed over pure secretions, their interest increasing and shifting over time, from sniffing and countermarking fresh mixtures, to licking and countermarking decayed mixtures. Substituting synthetic squalene (S)-a well-known fixative-for B secretions did not replicate prior results: B secretions, which contain additional chemicals that probably encode salient information, were preferred over pure S. Whereas support for the 'multiple-messages' hypothesis underscores the unique contribution from each of an animal's various secretions, support for the 'fixative' hypothesis highlights the synergistic benefits of composite signals.

Ca(2+) signals coordinate zygotic polarization and cell cycle progression in the brown alga Fucus serratus

Zygotes of the fucoid brown algae provide excellent models for addressing fundamental questions about zygotic symmetry breaking. Although the acquisition of polarity is tightly coordinated with the timing and orientation of the first asymmetric division-with zygotes having to pass through a G1/S-phase checkpoint before the polarization axis can be fixed -the mechanisms behind the interdependence of polarization and cell cycle progression remain unclear. In this study, we combine in vivo Ca(2+) imaging, single cell monitoring of S-phase progression and multivariate analysis of high-throughput intracellular Ca(2+) buffer loading to demonstrate that Ca(2+) signals coordinate polarization and cell cycle progression in the Fucus serratus zygote. Consistent with earlier studies on this organism, and in contrast to animal models, we observe no fast Ca(2+) wave following fertilization. Rather, we show distinct slow localized Ca(2+) elevations associated with both fertilization and S-phase progression, and we show that both S-phase and zygotic polarization are dependent on pre-S-phase Ca(2+) increases. Surprisingly, this Ca(2+) requirement cannot be explained by co-dependence on a single G1/ S-phase checkpoint, as S phase and zygotic polarization are differentially sensitive to pre-S-phase Ca(2+) elevations and can be uncoupled. Furthermore, subsequent cell cycle progression through M phase is independent of localized actin polymerization and zygotic polarization. This absence of a morphogenesis checkpoint, together with the observed Ca(2+)dependences of S phase and polarization, show that the regulation of zygotic division in the brown algae differs from that in other eukaryotic model systems, such as yeast and Drosophila.

Signals of need in a cooperatively breeding mammal with mobile offspring

In many bird species with biparental care for young in the nest, hungry chicks beg repeatedly and parents adjust their feeding rate to the call rate of young. Repetitive calling also occurs in fledglings and in some mammals where offspring follow provisioners. It is not yet clear whether, in mobile systems with dispersed young where adults cannot compare the vocal behaviour of all young simultaneously, the calls represent a signal of need. We investigated repetitive begging by cooperatively reared meerkat, Suricata suricatta, pups that foraged with the group. Pups produced two types of begging calls: repeat calls over long periods and high-pitched calls mainly confined to feeding events. Food-deprived pups stayed closer to feeders, and begged for longer and more intensely by calling at a higher rate. Hungry pups increased both the rate of repeat calls, which were given continually, and the number of high-pitched bouts, but adults increased their food allocation only in relation to the rate of repeat calls. Our study indicates that hunger may lead to several changes in vocal behaviour, only some of which may be used by adults to assess need.

Echolocation signals and pinnae movement in the fruitbat Rousettus aegyptiacus

The fruit bat Rousettus aegyptiacus has highly mobile pinnae. Little is known about the role that such movements play in sound localisation however and whether they interact with the process of echolocation in this species. Here we report the correspondence of echolocation signals in free flight with the downward wingbeat and forward movement of the pinnae, and demonstrate that the ears have a greater sensitivity to click stimuli in front of the animal when directed forwards than when back and to the side. The potential significance of the production of echolocation signals whilst the ears are moving from their least sensitive to their most sensitive position is discussed.

Convergent evolution of floral signals underlies the success of Neotropical orchids

The great majority of plant species in the tropics require animals to achieve pollination, but the exact role of floral signals in attraction of animal pollinators is often debated. Many plants provide a floral reward to attract a guild of pollinators, and it has been proposed that floral signals of non-rewarding species may converge on those of rewarding species to exploit the relationship of the latter with their pollinators. In the orchid family (Orchidaceae), pollination is almost universally animal-mediated, but a third of species provide no floral reward, which suggests that deceptive pollination mechanisms are prevalent. Here, we examine floral colour and shape convergence in Neotropical plant communities, focusing on certain food-deceptive Oncidiinae orchids (e.g. Trichocentrum ascendens and Oncidium nebulosum) and rewarding species of Malpighiaceae. We show that the species from these two distantly related families are often more similar in floral colour and shape than expected by chance and propose that a system of multifarious floral mimicry—a form of Batesian mimicry that involves multiple models and is more complex than a simple one model–one mimic system—operates in these orchids. The same mimetic pollination system has evolved at least 14 times within the species-rich Oncidiinae throughout the Neotropics. These results help explain the extraordinary diversification of Neotropical orchids and highlight the complexity of plant–animal interactions.