Influence of behavioural and morphological group composition on pigeon flocking dynamics.

Animals rely on movement to explore and exploit resources in their environment. While movement can provide energetic benefits, it also comes with energetic costs. This study examines how group phenotypic composition influences individual speed and energy expenditure during group travel in homing pigeons. We manipulated the composition of pigeon groups based on body mass and leadership rank. Our findings indicate that groups of 'leader' phenotypes show faster speeds and greater cohesion than 'follower' phenotype groups. Additionally, we show that groups of homogenous mass composition, whether all heavy or all light, were faster and expended less energy over the course of a whole flight than flocks composed of a mixture of heavy and light individuals. We highlight the importance of considering individual-level variation in social-level studies, and the interaction between individual and group-level traits in governing speed and the costs of travel.

'Selfish herders' finish last in mobile animal groups.

Predation is a powerful selective pressure and probably a driver of why many animal species live in groups. One key explanation for the evolution of sociality is the 'selfish herd' model, which describes how individuals who stay close to others effectively put neighbours between themselves and a predator to survive incoming attacks. This model is often illustrated with reference to herds of ungulates, schools of fish or flocks of birds. Yet in nature, when a predator strikes, herds are often found fleeing cohesively in the same direction, not jostling for position in the centre of the group. This paper highlights a critical assumption of the original model, namely that prey do not move in response to position of their predator. In this model, I relax this assumption and find that individuals who adopt 'selfish herd' behaviour are often more likely to be captured, because they end up at the back of a fleeing herd. By contrast, individuals that adopt a rule of 'neighbour to neighbour alignment' are able to avoid rearmost positions in a moving herd. Alignment is more successful than selfish herding across much of the parameter space, which may explain why highly aligned fleeing behaviour is commonly observed in nature.

Pigeon leadership hierarchies are not dependent on environmental contexts or individual phenotypes.

Remaining cohesive on the move can be beneficial for animal groups. As such, animal groups have evolved coordination mechanisms such as leadership to resolve navigational conflicts of interest. Consistent "leaders" may have an intrinsic advantage over "followers" which compromise on their preferred route to retain cohesion, which highlights the question of the inter-individual variation (phenotype) that can predict leadership. Studies in both birds and fish have revealed that intrinsically faster individuals can lead movements, and leading movements propagate from the front edge of the flock/shoal. However, these experiments are generally conducted in relatively "familiar" environments, where the degree of compromise between the "leaders" and "followers" is low. We suggested that inter-individual differences in route efficiency, while not explanatory of leadership from familiar locations, may emerge as predictors of leadership from unfamiliar locations. We tested this prediction - and the potential impact of multiple other behavioral, morphological and "in-flight" phenotypes on leadership - using two groups of homing pigeons (Columba livia) (N = 16), a classic model species of leadership. We recorded N = 966 unique GPS trajectories from birds in (i) solo and familiar, and (ii) solo and unfamiliar contexts to measure solo speed and solo route efficiency; and (iii) group and familiar, and (iv) group and unfamiliar contexts to assess group leadership. Pigeon leadership hierarchies were similar across environmental context (i.e., familiarity). However, we found that no covariates could consistently predict leadership score in either context.

Emergence of splits and collective turns in pigeon flocks under predation.

Complex patterns of collective behaviour may emerge through self-organization, from local interactions among individuals in a group. To understand what behavioural rules underlie these patterns, computational models are often necessary. These rules have not yet been systematically studied for bird flocks under predation. Here, we study airborne flocks of homing pigeons attacked by a robotic falcon, combining empirical data with a species-specific computational model of collective escape. By analysing GPS trajectories of flocking individuals, we identify two new patterns of collective escape: early splits and collective turns, occurring even at large distances from the predator. To examine their formation, we extend an agent-based model of pigeons with a 'discrete' escape manoeuvre by a single initiator, namely a sudden turn interrupting the continuous coordinated motion of the group. Both splits and collective turns emerge from this rule. Their relative frequency depends on the angular velocity and position of the initiator in the flock: sharp turns by individuals at the periphery lead to more splits than collective turns. We confirm this association in the empirical data. Our study highlights the importance of discrete and uncoordinated manoeuvres in the collective escape of bird flocks and advocates the systematic study of their patterns across species.

Self-organization of collective escape in pigeon flocks.

Bird flocks under predation demonstrate complex patterns of collective escape. These patterns may emerge by self-organization from local interactions among group-members. Computational models have been shown to be valuable for identifying what behavioral rules may govern such interactions among individuals during collective motion. However, our knowledge of such rules for collective escape is limited by the lack of quantitative data on bird flocks under predation in the field. In the present study, we analyze the first GPS trajectories of pigeons in airborne flocks attacked by a robotic falcon in order to build a species-specific model of collective escape. We use our model to examine a recently identified distance-dependent pattern of collective behavior: the closer the prey is to the predator, the higher the frequency with which flock members turn away from it. We first extract from the empirical data of pigeon flocks the characteristics of their shape and internal structure (bearing angle and distance to nearest neighbors). Combining these with information on their coordination from the literature, we build an agent-based model adjusted to pigeons' collective escape. We show that the pattern of turning away from the predator with increased frequency when the predator is closer arises without prey prioritizing escape when the predator is near. Instead, it emerges through self-organization from a behavioral rule to avoid the predator independently of their distance to it. During this self-organization process, we show how flock members increase their consensus over which direction to escape and turn collectively as the predator gets closer. Our results suggest that coordination among flock members, combined with simple escape rules, reduces the cognitive costs of tracking the predator while flocking. Such escape rules that are independent of the distance to the predator can now be investigated in other species. Our study showcases the important role of computational models in the interpretation of empirical findings of collective behavior.

Absence of "selfish herd" dynamics in bird flocks under threat.

The "selfish herd" hypothesis1 provides a potential mechanism to explain a ubiquitous phenomenon in nature: that of non-kin aggregations. Individuals in selfish herds are thought to benefit by reducing their own risk at the expense of conspecifics by attracting toward their neighbors' positions1,2 or central locations in the aggregation.3-5 Alternatively, increased alignment with their neighbors' orientation could reduce the chance of predation through information sharing6-8 or collective escape.6 Using both small and large flocks of homing pigeons (Columba livia; n = 8-10 or n = 27-34 individuals) tagged with 5-Hz GPS loggers and a GPS-tagged, remote-controlled model peregrine falcon (Falco peregrinus), we tested whether individuals increase their use of attraction over alignment when under perceived threat. We conducted n = 27 flights in treatment conditions, chased by the robotic "predator," and n = 16 flights in control conditions (not chased). Despite responding strongly to the RobotFalcon-by turning away from its flight direction-individuals in treatment flocks demonstrated no increased attraction compared with control flocks, and this result held across both flock sizes. We suggest that mutualistic alignment is more advantageous than selfish attraction in groups with a high coincidence of individual and collective interests (adaptive hypothesis). However, we also explore alternative explanations, such as high cognitive demand under threat and collision avoidance (mechanistic hypotheses). We conclude that selfish herd may not be an appropriate paradigm for understanding the function of highly synchronous collective motion, as observed in bird flocks and perhaps also fish shoals and highly aligned mammal aggregations, such as moving herds.

Artificial mass loading disrupts stable social order in pigeon dominance hierarchies.

Dominance hierarchies confer benefits to group members by decreasing the incidences of physical conflict, but may result in certain lower ranked individuals consistently missing out on access to resources. Here, we report a linear dominance hierarchy remaining stable over time in a closed population of birds. We show that this stability can be disrupted, however, by the artificial mass loading of birds that typically comprise the bottom 50% of the hierarchy. Mass loading causes these low-ranked birds to immediately become more aggressive and rise-up the dominance hierarchy; however, this effect was only evident in males and was absent in females. Removal of the artificial mass causes the hierarchy to return to its previous structure. This interruption of a stable hierarchy implies a strong direct link between body mass and social behaviour and suggests that an individual's personality can be altered by the artificial manipulation of body mass.

When flocking is costly: reduced cluster-flock density over long-duration flight in pigeons.

Birds which fly in coordinated cluster-flocks can benefit through the formation of group-level structures and patterns which can deter predators by visual confusion. Though unlike V-formation flight, cluster-flocking increases the energetic cost of flight, particularly in denser flocks. Cluster formations therefore provide a unique opportunity to investigate trade-offs between increased work rate (e.g. higher flap frequency) and other benefits of flocking. As part of a routine 9-km training flight release, a flock of six homing pigeons (Columba livia) with 5 Hz GPS and 200 Hz accelerometer biologgers attached flew an alternative trajectory totalling 177 km and 256 min of flight. We provide the first evidence that during a long-duration flight, pigeons' pairwise and group-level distances increased (i.e. group structure changed), while flap frequency decreased over time. This implies that as birds tire during long-duration flight, the ultimate functions of cluster-flocking-primarily anti-predator benefits-are overridden by the proximate costs of flying close to conspecifics.

Lensless phase contrast microscopy based on multiwavelength Fresnel diffraction.

We demonstrate a compact, wide-field, quantitative phase contrast microscope that does not require lenses for image formation. High-resolution images are retrieved from Fresnel diffraction patterns recorded at multiple wavelengths, combined with a robust iterative phase retrieval algorithm. Quantitative phase contrast images of living cultured neurons are obtained with a transverse resolution of <2 µm. Our system is well suited for high-resolution live cell imaging and provides a compact, cost-effective alternative to full-sized phase-contrast microscopes.

High-speed multi-wavelength Fresnel diffraction imaging.

We demonstrate a compact lensless microscope which can capture video-rate phase contrast images of moving objects and allows numerical scanning of the focal distance after recording. Using only an RGB-detector and illumination from a single mode fiber, diffraction patterns at three wavelengths are recorded simultaneously, enabling high-speed data collection and reconstruction of phase and amplitude. The technique is used for imaging of a moving test target, beads in a flow cell, and imaging of Caenorhabditis elegans moving in a droplet of liquid.

High-energy, high-repetition-rate picosecond pulses from a quasi-CW diode-pumped Nd:YAG system.

We report on a high-power quasi-CW pumped Nd:YAG laser system, producing 130 mJ, 64 ps pulses at 1064 nm wavelength with a repetition rate of 300 Hz. Pulses from a Nd:YVO(4) oscillator are first amplified by a regenerative amplifier to the millijoule level and then further amplified in quasi-CW diode-pumped Nd:YAG modules. Pulsed diode pumping enables a high gain at repetition rates of several hundred hertz, while keeping thermal effects manageable. Birefringence compensation and multiple thermal-lensing-compensated relay-imaging stages are used to maintain a top-hat beam profile. After frequency doubling, 75 mJ pulses are obtained at 532 nm. The intensity stability is better than 1.1%, which makes this laser an attractive pump source for a high-repetition-rate optical parametric amplification system.

Electron beam intraoperative radiation therapy for pediatric neoplasms.

BACKGROUND: Intraoperative radiation therapy (IORT) has potential advantages over conventional external beam radiation in that a single large dose is delivered to the tumor and its regional bed at the time of surgical exploration. The therapeutic ratio is enhanced by direct tumor visualization, precise treatment volume, and exclusion of normal organs. In childhood, local tumor control is critical for lesions that cannot be totally excised or residual disease not ablated by systemic therapy. METHODS: During the past decade, the authors evaluated IORT in pediatric patients with unresectable or recurrent tumors. Fifty-nine patients were treated, 48 for advanced malignant disease and 11 for histologically benign but locally aggressive tumors. Sixty-four operations were performed, during which 84 separate radiation fields were used. High energy electrons at 5-11 MeV were delivered at a dose of 1000-1700 cGy to a tissue depth of 0.5-3 cm. RESULTS: Of 25 children with advanced neuroblastoma, 15 were alive 14-104 months (mean, 51 months) after treatment. The survival of all patients with solid malignancies was 63%. Local tumor control was achieved in 75% of children with cancer and 91% of those with benign tumors. Complications of IORT at the doses used were trivial. CONCLUSIONS: Electron beam therapy can be safely and efficiently delivered to young patients during operations intended to treat the primary tumor. The outcome achieved for overall survival and local control of the primary tumor is encouraging, and an acceptable complication rate during intermediate to long-term follow-up has been noted.

AIDS-related non-Hodgkin's lymphoma: the outcome and efficacy of radiation therapy.

The records of all 16 patients with AIDS-related lymphoma treated with radiation therapy at our institutions were reviewed. All patients were male with a median age of 32 years, and all but one had biopsy proven high-grade NHL. Eleven had lymphoma involving the central nervous system and five had lymphoma involving other sites. Seven of the 11 patients with CNS involvement had primary CNS lymphoma. All patients were treated with megavoltage X rays to doses ranging from 1050 cGy in 1 1/2 weeks to 5037 cGy in 6 weeks. Of those patients with CNS lymphoma, only one responded completely and four responded partially to irradiation. All patients died within a range of 0.2 to 5.3 months (median survival = 2.2 months) from starting radiation therapy. In contrast, 3 of 5 patients (60%) with NHL outside the CNS responded completely and 1 responded partially to involved-field irradiation. These patients survived a median of 12.6 months with one achieving long-term lymphoma-free survival at 40 months. This long-term survivor presented with Stage IE lymphoma as his only manifestation of AIDS. We conclude that AIDS-related lymphomas respond less favorably to radiation therapy than lymphomas in non-immunosuppressed patients. Furthermore, CNS lymphomatous involvement is an ominous occurrence in the AIDS patient. In our experience, cranial irradiation failed to provide significant palliation or survival prolongation in this group of patients. Instead, long-term survival is possible in AIDS patients with limited NHL outside the CNS, and it is in these patients that combination chemotherapy plus involved-field radiation therapy may play a curative role.

Anesthesia for intraoperative radiation therapy in children.

Intraoperative radiation therapy (IORT) is a relatively new mode of cancer treatment which is being used with increasing frequency. IORT presents several challenges to the anesthesiologist, including patients who are debilitated from their disease or chemotherapy, operations involving major tumor resections, intraoperative interdepartmental transport of patients, and remote monitoring of patients during electron beam therapy. This report discusses the anesthetic management of ten children undergoing IORT. With adequate preparation and interdepartmental communication, complications can be avoided during these challenging cases.

Follow-up usage of the scott-craig Orthosis in paraplegia.

A survey of the usage of the Scott-Craig Orthosis among individuals with LI or above complete motor and sensory paraplegia is reported. Patients were grouped according to those without volitionally controlled abdominal musculature (C8-T5), those with volitionally controlled upper abdominal musculature (T6-T9), and those with all abdominal musculature under volitional control (T10-LI). The study was done in two phases. Phase I, done in 1976, consisted of a survey of 184 patients in this neurological grouping who received the Scott-craig Orthosis at Craig Hospital from 1 January 1964 through to 31 December 1975. Phase II was completed in December 1980. Sixty-three additional patients were surveyed who were braced from 1 January 1976 through to 31 December 1979. Long-term usage and some form of ambulation was found in 77 per cent of those surveyed.

Spin-labeled ribonuclease A. Effects of chemical, enzymatic, and physical modifications on enzyme conformation.

3-SLHis-105-RNase A is an active derivative of ribonuclease A (RNase A) spin-labeled at the 3 position of the imidazole ring of histidine-105. The spin-labeled enzyme has been modified by urea denaturation, reduction, reduction-carboxymethylation, performic acid oxidation, and digestion with proteolytic enzymes in order to monitor changes in the geometry of the protein by changes in the electron paramagnetic resonance (EPR) spectrum of the nitroxide spin-label probe. The results of these experiments indicate that the spin-label attached to histidine-105 of RNase A is sensitive to modifications affecting the conformational integrity of the molecule and to the reconstituting effects of various active-center ligands.

19F nuclear magnetic resonance of 5-fluorouridine-substituted tRNA1Val from Escherichia coli.

The 19F NMR spectrum of Escherichia coli tRNA1Val in which [5-19F]uridine replaces 93% of all uridine and uridine-derived residues has been examined at 93.6 and 235 MHz. The resolution of 11 peaks and visibility of two additional shoulders at either frequency for the 14 FUra residues in the molecule attests to the excellence of 19F as a probe for the structure of tRNA1Val in solution. No significant gain in resolution was attained at the higher frequency. A comparison of the relative areas in the different regions of the 19F spectrum of mixed [FUra]tRNAs with that of [FUra]tRNA1Val suggests that the three single resonances at lowest field in the region 86.5 to 88.5 ppm upfield from trifluoroacetate correspond to the three invariant bases which form tertiary hydrogen bonds in all tRNAs, namely, 8 (U or s4U), 54 (T), and 55 (phi) in unsubstituted tRNAs.

lac repressor: 3-fluorotyrosine substitution for nuclear magnetic resonance studies.

This paper describes the isolation of 3-fluorotyrosine-substituted lac repressor, and its 19F nuclear magnetic resonance spectrum. From the spectrum, one can conclude that for each of the four identical subunits of the repressor there are four or five surface tyrosines, two buried or internal tyrosines, and one tyrosine with an phenolic group ionized or involved in a hydrogen bond. Conditions are described that can be used for the 3-fluorotyrosine substitution of a variety of Escherichia coli proteins for 19F nuclear magnetic resonance studies.

Changes in tertiary structure accompanying a single base change in transfer RNA. Proton magnetic resonance and aminoacylation studies of Escherichia coli tRNAMet f1 and tRNAMet f3 and their spin-labeled (s4U8) derivatives.

The properties of Escherichia coli tRNAMet f1 and tRNAMet f3 that differ by only one base change, m7G to A at position 47, have been compared structurally by proton magnetic resonance and functionally by the aminoacylation reaction. The NMR spectra of the two tRNA species in the region between 0 and 4 ppm below 4,4-dimethyl-4-silapentane-1-sulfonic acid (DSS) (methyl and methylene region) were the same except for the absence of the lowest field peak at 3.8 ppm in tRNAMet f3, thus unequivocally identifying this resonance at the methyl group of m7G47 of tRNAMet f1. The same resonance disappears in tRNAMet f1 spin-labeled at s4U8 and reappears in the diamagnetic reduced spin-labeled tRNAMet f1 from which the average distance between the spin-label and the methyl protons of m7G is estimated to be less than 15 A. The proximity of m7G47 but not T55 to s4U8 in the structure of E. coli tRNAMet f1 in solution is consistant with the crystallographic model for yeast tRNAPhe. A spectral comparison of the hydrogen-bond regions (11-14 ppm below DSS) of tRNAMet f1 and tRNAMet f3 reveals major shifts of four resonances previously assigned to tertiary hydrogen bonds. Of the four, the one at lowest field (14.8 ppm) had been assigned by chemical modification to the tertiary (s4U8-A14) hydrogen bond and the one at 13.3 ppm had been tentatively assigned to the tertiary hydrogen bond G23-m7G47 of the 13-23-47 triple. A more positive assignment of the G23-m7G47 at 13.3 ppm could be made from the additional evidence that this resonance, which was first observed in the difference spectrum between spin-labeled tRNAMet f1 and its reduced form, is the only one missing in the analogous difference spectrum of tRNAMet f3. At low ionic strength and in the absence of magnesium ions, the differences in the hydrogen-bonded region of the NMR spectra of tRNAMet f1 and tRNAMet f3 are much greater than in the presence of magnesium ions. The optimal magnesium concentration required for maximal initial velocities is also higher for tRNAMet f3 than for tRNAMet f1. The perturbation caused by the spin-label in destabilizing hydrogen bonds in the region between 13 and 14 ppm is greater for tRNAMet f3 than tRNAMet f1 but the distance relations for the hydrogen bonds in the region between 12 and 13 ppm (the major paramagnetic perturbations) are conserved in the two species. The disruption of one hydrogen bond relative to native tRNAMet f1 either by spin-labeling (s4U8-A14) or by substitution of m7G by A in tRNAMet f3 has little effect on the aminoacyl acceptor activity or the velocity of the aminoacylation reaction at optimal magnesium concentration, but the absence of both tertiary hydrogen bonds in the augmented D-helix region in the spin-labeled tRNAMet f3 results in approximately 60% reduction both in acceptance activity and in initial velocity of the aminoacylation reaction.

Nuclear magnetic resonance study of hydrogen-bonded ring protons in oligonucleotide helices involving classical and nonclassical base pairs.

A study of the exchangeable ring nitrogen protons in aqueous solutions of oligonucleotide complexes involving Watson-Crick base pairs as well as Hoogsteen pairs and other nonclassical hydrogen bonding schemes shows that resolvable resonances in the low-field (-10 to -16 ppm from sodium 4,4-dimethyl-4-silapentanesulfonate) region can be detected in a variety of structures other than double stranded helices. Ring nitrogen proton resonances arising from the following hydrogen-bonding situations are reported: (1) AT and GC Watson-Crick base pairs in a self-complementary octanucleotide, dApApApGpCpTpTpT; (2) U-A-U base triples in complexes between oligo-U15 and AMP; (3) C-G-C+ base triples in complexes between oligo-C17 and GMP at acid pH; (4) s4U-A-s4U base triples in complexes between oligo-s4U15 and AMP, all of which involve both Watson-Crick and Hoogsteen base pairing to form triplexes; (5) C-C+ base pairing between protonated and unprotonated C residues in oligo-C17 at acid pH; and (6) I4 base quadruples in the four strand association among oligo-I at high salt. The behavior of the dA3G-CT3 helix is consistent with both fraying of the terminal base pairs and presence of intermediate states as the helix opens. In the monomer-oligomer complexes, under the conditions used here, the exchange appears to be governed by the dissociation rate of monomer from the complex. These findings suggest that those tertiary structure hydrogen bonds in tRNA involving ring nitrogen protons should have representative resonances in the low-field (11-16 ppm) proton NMR region in H2O.