High-Q nanomechanics via destructive interference of elastic waves.

Mechanical dissipation poses a ubiquitous challenge to the performance of nanomechanical devices. Here we analyze the support-induced dissipation of high-stress nanomechanical resonators. We develop a model for this loss mechanism and test it on Si(3)N(4) membranes with circular and square geometries. The measured Q values of different harmonics present a nonmonotonic behavior which is successfully explained. For azimuthal harmonics of the circular geometry we predict that destructive interference of the radiated waves leads to an exponential suppression of the clamping loss in the harmonic index. Our model can also be applied to graphene drums under high tension.

Terahertz imaging and spectroscopy of large-area single-layer graphene.

We demonstrate terahertz (THz) imaging and spectroscopy of a 15 × 15-mm2 single-layer graphene film on Si using broadband THz pulses. The THz images clearly map out the THz carrier dynamics of the graphene-on-Si sample, allowing us to measure sheet conductivity with sub-mm resolution without fabricating electrodes. The THz carrier dynamics are dominated by intraband transitions and the THz-induced electron motion is characterized by a flat spectral response. A theoretical analysis based on the Fresnel coefficients for a metallic thin film shows that the local sheet conductivity varies across the sample from σ(s) = 1.7 × 10(-3) to 2.4 × 10(-3) Ω(-1) (sheet resistance, ρ(s) = 420 - 590 Ω/sq).

Real-time synchronous imaging of electromechanical resonator mode and equilibrium profiles.

Interferometric imaging of normal mode dynamics in electromechanical resonators, oscillating in the rf regime, is demonstrated by synchronous imaging with a pulsed nanosecond laser. Profiles of mechanical modes in suspended thin film structures and their equilibrium profiles are measured through all-optical Fabry-Perot reflectance fits to the temporal traces. As a proof of principle, the mode patterns of a microdrum silicon resonator are visualized, and the extracted vibration modes and equilibrium profile show good agreement with numerical estimations.

Stress and silicon nitride: a crack in the universal dissipation of glasses.

High-stress silicon nitride microresonators exhibit a remarkable room temperature Q factor that even exceeds that of single crystal silicon. A study of the temperature dependent variation of the Q of a 255 micromx255 micromx30 nm thick high-stress Si3N4 membrane reveals that the dissipation Q-1 decreases with lower temperatures and is approximately 3 orders of magnitude smaller than the universal behavior. Stress-relieved cantilevers fabricated from the same material show a Q that is more consistent with typical disordered materials. e-beam and x-ray studies of the nitride film's structure reveal characteristics consistent with a disordered state. Thus, it is shown that stress alters the Q-1, violating the universality of dissipation in disordered materials in a self-supporting structure.

Sperm competition and brain size evolution in mammals.

The 'expensive tissue hypothesis' predicts a size trade-off between the brain and other energetically costly organs. A specific version of this hypothesis, the 'expensive sexual tissue hypothesis', argues that selection for larger testes under sperm competition constrains brain size evolution. We show here that there is no general evolutionary trade-off between brain and testis mass in mammals. The predicted negative relationship between these traits is not found for rodents, ungulates, primates, carnivores, or across combined mammalian orders, and neither does total brain mass vary according to the level of sperm competition as determined by mating system classifications. Although we are able to confirm previous reports of a negative relationship between brain and testis mass in echolocating bats, our results suggest that mating system may be a better predictor of brain size in this group. We conclude that the expensive sexual tissue hypothesis accounts for little or none of the variance in brain size in mammals, and suggest that a broader framework is required to understand the costs of brain size evolution and how these are met.

Energetic constraints, not predation, influence the evolution of sleep patterning in mammals.

Mammalian sleep is composed of two distinct states - rapid-eye-movement (REM) and non-REM (NREM) sleep - that alternate in cycles over a sleep bout. The duration of these cycles varies extensively across mammalian species. Because the end of a sleep cycle is often followed by brief arousals to waking, a shorter sleep cycle has been proposed to function as an anti-predator strategy. Similarly, higher predation risk could explain why many species exhibit a polyphasic sleep pattern (division of sleep into several bouts per day), as having multiple sleep bouts avoids long periods of unconsciousness, potentially reducing vulnerability.Using phylogenetic comparative methods, we tested these predictions in mammals, and also investigated the relationships among sleep phasing, sleep-cycle length, sleep durations and body mass.Neither sleep-cycle length nor phasing of sleep was significantly associated with three different measures of predation risk, undermining the idea that they represent anti-predator adaptations.Polyphasic sleep was associated with small body size, shorter sleep cycles and longer sleep durations. The correlation with size may reflect energetic constraints: small animals need to feed more frequently, preventing them from consolidating sleep into a single bout. The reduced daily sleep quotas in monophasic species suggests that the consolidation of sleep into one bout per day may deliver the benefits of sleep more efficiently and, since early mammals were small-bodied and polyphasic, a more efficient monophasic sleep pattern could be a hitherto unrecognized advantage of larger size.

Evolutionary specialization in mammalian cortical structure.

Changes in neocortex size were a prominent feature of mammalian brain evolution, but the implications for cortical structure, and consequently for the functional significance of such changes in overall cortical size, are poorly understood. A basic question is whether functionally differentiated cortical areas evolved independently of one another (adaptive specialization) or were allometrically constrained to co-vary tightly with the size of the whole. Here, I provide comparative evidence for adaptive specialization of cortical structure. First, the sizes of individual areas differ significantly between taxa after controlling for overall cortical size. Second, an analysis of separate visual cortical areas reveals that these exhibit statistically correlated evolution, independent of variation in nonvisual areas. Third, visual cortex size exhibits correlated evolution with peripheral visual adaptations (eye morphology and optic nerve size) and with photic niche. Thus, the evolution of mammalian cortical structure was closely associated with specialization for different sensory niches.

From The Cover: Binocularity and brain evolution in primates.

Primates are distinguished by frontally directed, highly convergent orbits, which are associated with stereoscopic vision. Although stereoscopic vision requires specialized neural mechanisms, its implications for brain evolution are unknown. Using phylogenetic comparative analysis, I show that evolutionary increases among primate taxa in the degree of orbital convergence correlate with expansion of visual brain structures and, as a consequence, with the overall size of the brain. This pattern is found across the whole primate order and is also repeated within each of the two major primate subtaxa. The visual expansion associated with increased binocularity is specific to the parvocellular visual pathway, consistent with recent evidence implicating this pathway in fine-grained stereopsis. The results support the hypothesis that brain size evolution in primates was associated with visual specialization.

The evolution of the cortico-cerebellar complex in primates: anatomical connections predict patterns of correlated evolution.

Investigations into the evolution of the primate brain have tended to neglect the role of connectivity in determining which brain structures have changed in size, focusing instead on changes in the size of the whole brain or of individual brain structures, such as the neocortex, in isolation. We show that the primate cerebellum, neocortex, vestibular nuclei and relays between them exhibit correlated volumetric evolution, even after removing the effects of change in other structures. The patterns of correlated evolution among individual nuclei correspond to their known patterns of connectivity. These results support the idea that the brain evolved by mosaic size change in arrays of functionally connected structures. Furthermore, they suggest that the much discussed expansion of the primate neocortex should be re-evaluated in the light of conjoint cerebellar expansion.

Event-by-event fluctuations of the Kaon-to-Pion ratio in central Pb+Pb collisions at 158 GeV per nucleon.

We present the first measurement of fluctuations from event to event in the production of strange particles in collisions of heavy nuclei. The ratio of charged kaons to charged pions is determined for individual central Pb+Pb collisions. After accounting for the fluctuations due to detector resolution and finite number statistics we derive an upper limit on genuine nonstatistical fluctuations, which could be related to a first- or second-order QCD phase transition. Such fluctuations are shown to be very small.

Mosaic evolution of brain structure in mammals.

The mammalian brain comprises a number of functionally distinct systems. It might therefore be expected that natural selection on particular behavioural capacities would have caused size changes selectively, in the systems mediating those capacities. It has been claimed, however, that developmental constraints limited such mosaic evolution, causing co-ordinated size change among individual brain components. Here we analyse comparative data to demonstrate that mosaic change has been an important factor in brain structure evolution. First, the neocortex shows about a fivefold difference in volume between primates and insectivores even after accounting for its scaling relationship with the rest of the brain. Second, brain structures with major anatomical and functional links evolved together independently of evolutionary change in other structures. This is true at the level of both basic brain subdivisions and more fine-grained functional systems. Hence, brain evolution in these groups involved complex relationships among individual brain components.

Visual specialization and brain evolution in primates.

Several theories have been proposed to explain the evolution of species differences in brain size, but no consensus has emerged. One unresolved question is whether brain size differences are a result of neural specializations or of biological constraints affecting the whole brain. Here I show that, among primates, brain size variation is associated with visual specialization. Primates with large brains for their body size have relatively expanded visual brain areas, including the primary visual cortex and lateral geniculate nucleus. Within the visual system, it is, in particular, one functionally specialized pathway upon which selection has acted: evolutionary changes in the number of neurons in parvocellular, but not magnocellular, layers of the lateral geniculate nucleus are correlated with changes in both brain size and ecological variables (diet and social group size). Given the known functions of the parvocellular pathway, these results suggest that the relatively large brains of frugivorous species are products of selection on the ability to perceive and select fruits using specific visual cues such as colour. The separate correlation between group size and visual brain evolution, on the other hand, may indicate the visual basis of social information processing in the primate brain.

Neocortex size and behavioural ecology in primates.

The neocortex is widely held to have been the focus of mammalian brain evolution, but what selection pressures explain the observed diversity in its size and structure? Among primates, comparative studies suggest that neocortical evolution is related to the cognitive demands of sociality, and here I confirm that neocortex size and social group size are positively correlated once phylogenetic associations and overall brain size are taken into account. This association holds within haplorhine but not strepsirhine primates. In addition, the neocortex is larger in diurnal than in nocturnal primates, and among diurnal haplorhines its size is positively correlated with the degree of frugivory. These ecological correlates reflect the diverse sensory-cognitive functions of the neocortex.

Evolutionary radiation of visual and olfactory brain systems in primates, bats and insectivores.

How brains have evolved in response to particular selection pressures is illuminated by ecological correlates of differences in brain structure among contemporary species. The focus of most comparative studies has been on the overall size of brains relative to body size, hence ignoring the ways in which selection operates on specific neural systems. Here we investigate evolutionary radiations in the size of visual and olfactory brain structures within three orders of mammals: primates, bats and insectivores. The comparative relationships within these three orders show both similarities and differences. After removal of the allometric effect of overall brain size, the sizes of different structures within each sensory modality are positively correlated in all three orders. Correlations between visual and olfactory structures, however, are negative in primates, negative but non-significant in insectivores, and positive in bats. In both primates and insectivores, nocturnal lineages tend to have larger olfactory structures than do diurnal or partly diurnal lineages, and among the primates diurnal lineages have larger striate visual cortexes. Hence the apparent trade-off between vision and olfaction in primates seems to be related to the divergence of nocturnal and diurnal forms. However, negative correlations between visual and olfactory structures were also found when nocturnal strepsirhines and diurnal haplorhines were analysed separately, suggesting that ecological variables in addition to activity timing may be significant. Indeed, there were also associations with diet: frugivory was associated with enlargements of the geniculostriate visual system in diurnal primates, enlargements of olfactory structures in nocturnal primates, and possibly enlargements of both in bats. Further ecological associations were found within insectivores: aquatic lineages had smaller olfactory structures than in their non-aquatic counterparts, and fossorial lineages had smaller optic nerves than in non-fossorial forms. We conclude that activity timing, diet and habitat have each played a role in the evolutionary radiation of mammalian sensory systems, but with varying effects in the different taxa. Some of the associations between ecology and sensory systems suggest alternative explanations for correlates of overall brain size, which have in the past commonly been interpreted in terms of selection on intelligence.

Comparative evidence indicating neural specialization for predatory behaviour in mammals.

The evolution of cognitive and sensory specializations must involve concomitant modifications of neural substrates. Ecological correlates of species differences in brain structure are intriguing sources of evidence about such evolutionary specialization but, to date, these have been identified only for gross parameters, such as overall brain size and the size of major brain regions. Here we show that a behavioural specialization in mammals, predation, is associated with species differences in the fine structure of a single neural pathway, the tectospinal tract. Both the relative number of neurons in this pathway and the relative size of their cell bodies were greater in more predatory species than in their less predatory counterparts within each of four separate mammalian orders. Expansion of these analyses to consider comparisons between taxa at a variety of taxonomic levels gave further support to the idea of a relation between predatory habits and the evolution of the tectospinal tract. In addition, within the primates, the number of neurons in the tectospinal tract was significantly correlated with the proportion of prey in the diet. These results therefore appear to provide an example of correlated evolution between a specific neural system and behaviour which applies generally within the mammals. They also help to unify findings from physiological and anatomical studies on a wider range of vertebrate taxa, including reptiles and amphibians.

Chemical composition of baboon plant foods: implications for the interpretation of intra- and interspecific differences in diet.

Information on the chemical composition of baboon foods from the Laikipia Plateau, Kenya, is presented. Despite some differences in methods, results of analyses performed on the same foods at different sites were found to be extremely consistent, encouraging the view that meaningful intra- and interspecific comparisons of diet selection are feasible. Contrary to assumptions in the literature, no relationship between the abundance of food types and their chemical composition was found, nor was the foliage eaten by the baboons found to be a low-quality or high-fibre item in comparison with fruits and storage organs. Emphasis is placed on the need for caution in the use of simplistic dietary taxonomies which imply phytochemical and ecological homogeneity within broad food categories. Comparisons between three species revealed marked differences in the chemical composition of their diets; in particular, baboon diets were found to be higher in protein and lower in fibre than those of either lowland gorillas or Malaysian leaf monkeys, and differences in condensed tannin levels were also found. The relationship between these differences and the socio-ecology of the three species is discussed.

Meat quality and muscle fibre type characteristics of Southdown Rams from high and low backfat selection lines.

Characteristics of the meat of 15-18-month Southdown rams from lines selected for high or low backfat depths (assessed ultrasonically at position C over the last rib) were compared. Half of the carcasses were electrically stimulated (ES) and within each carcass post-mortem treatments chosen to produce effects on meat tenderness were ageing periods of 1 or 15 days (Semimembranosus), early or delayed chilling (Biceps femoris), and trimming of the s.c. fat cover (Longissimus dorsi). These treatments had the expected effects on shear values, but the sizes of the effects were little affected by selection line or ES treatment. Selection line did not have any direct effects on shear values, reflectance values at several wavelengths, waterholding capacity, cooking loss or sarcomere length. The Semitendinosus muscle had a higher proportion of predominantly oxidative fibres for the high-backfat line, based on succinic dehydrogenase activity (P < 0·05), but there was no line difference in alkaline-stable ATPase activity in the same muscle. Muscle fibre diameter was similar for the two lines.

Dietary and foraging strategies of baboons.

As large-bodied savannah primates, baboons have long been of special interest to students of human evolution: many different populations have been studied and dietary comparisons among them are becoming possible. Baboons' foraging strategies can be shown to combine high degrees of flexibility and breadth with selectivity. In this paper we develop and test multivariate models of the basis of diet selection for populations of montane and savannah baboons. Food selection is positively related to protein and lipid content and negatively to fibre, phenolics and alkaloids. Seasonal changes in dietary criteria predicted by these rules are tested and confirmed. Although nutritional bottlenecks occur at intervals, a comparison between long-term nutrient intakes in four different populations indicates convergence on lower degrees of variation than exist in superficial foodstuff profiles.

Demise of the checksheet: Using off-the-shelf miniature hand-held computers for remote fieldwork applications.

Laboratory-based researchers have increasingly reaped the benefits of entering data directly into a computer; those concerned with behaviour often using specially designed keyboards. However, many ecologists and ethologists doing fieldwork in remote places have been reluctant to abandon paper checksheets because of worries about unreliability, lack of electrical supply and sheer weight of computer equipment, adding to more general drawbacks such as the need for considerable expertise in purpose-built hardware and software. Having used commercially available hand-held computers extensively for our own fieldwork on baboons in Africa, we are confident that these worries are unfounded. As some researchers have already discovered, field computerization is not something to be distrusted, but in fact offers several important benefits.