Impact Loading and Locomotor-Respiratory Coordination Significantly Influence Breathing Dynamics in Running Humans

Locomotor-respiratory coupling (LRC), phase-locking between breathing and stepping rhythms, occurs in many vertebrates. When quadrupedal mammals gallop, 1∶1 stride per breath coupling is necessitated by pronounced mechanical interactions between locomotion and ventilation. Humans show more flexibility in breathing patterns during locomotion, using LRC ratios of 2∶1, 2.5∶1, 3∶1, or 4∶1 and sometimes no coupling. Previous studies provide conflicting evidence on the mechanical significance of LRC in running humans. Some studies suggest LRC improves breathing efficiency, but others suggest LRC is mechanically insignificant because ‘step-driven flows’ (ventilatory flows attributable to step-induced forces) contribute a negligible fraction of tidal volume. Yet, although step-driven flows are brief, they cause large fluctuations in ventilatory flow. Here we test the hypothesis that running humans use LRC to minimize antagonistic effects of step-driven flows on breathing. We measured locomotor-ventilatory dynamics in 14 subjects running at a self-selected speed (2.6±0.1 ms⁻¹) and compared breathing dynamics in their naturally ‘preferred’ and ‘avoided’ entrainment patterns. Step-driven flows occurred at 1-2X step frequency with peak magnitudes of 0.97±0.45 Ls⁻¹ (mean ±S.D). Step-driven flows varied depending on ventilatory state (high versus low lung volume), suggesting state-dependent changes in compliance and damping of thoraco-abdominal tissues. Subjects naturally preferred LRC patterns that minimized antagonistic interactions and aligned ventilatory transitions with assistive phases of the step. Ventilatory transitions initiated in ‘preferred’ phases within the step cycle occurred 2x faster than those in ‘avoided’ phases. We hypothesize that humans coordinate breathing and locomotion to minimize antagonistic loading of respiratory muscles, reduce work of breathing and minimize rate of fatigue. Future work could address the potential consequences of locomotor-ventilatory interactions for elite endurance athletes and individuals who are overweight or obese, populations in which respiratory muscle fatigue can be limiting.     

Axial-Appendicular Dynamics and the Integration of Breathing and Gait in Mammal

SYNOPSIS. This paper proposes a biomechanical model for locomotor-respiratorycoupling (LRC) in galloping mammals in which gait and breathingcycles are phase-locked on a 1:1 basis. It also explores someof the physiological and neuromotor implications of LRC. The mechanical coupling of locomotor and respiratory cyclesdepends upon the coordinated, reciprocal oscillations of thecranio-cervical and lumbo-pelvic components of the axial systemand their attendant actions on the intervening thorax via muscularlinkages. Concurrently, accelerational and decelerational forcesimparted to the axial system by the limbs help to drive lungventilation by inducing inertial displacements of a "visceralpiston’ connected to the diaphragm. Several lines of evidence(including cineradiographic data) suggest that an importantfunction of the crural diaphragm is to control the displacementof the visceral piston. The kinematics of LRC indicate thatthe interosseous intercostal muscles must simultaneously operateto assure thoracic stability against locomotor stresses as wellas to promote breathing. The former may be their more essentialrole, however. The characteristic design of the rib cage incursorial mammals (=deep and narrow) appears to maximize theleverage of certain "accessory respiratory muscles" (i.e., sternocleidomastoid,scalenes) while minimizing torsional loading of the thorax duringforelimb support. Physiological implications of LRC include the prediction thatlarge mammals will breathe relatively faster and with relativelysmaller lung volumes when galloping than small species. An additionalprediction, that running mammals could automatically gear lungventilation to speed by simply linking breathing rate to stridefrequency and depth of breath (=tidal volume) to stride length,appears to be supported by experimental data from horses. Finally,the neuromotor basis of LRC probably depends upon the directinteraction of central pattern generators for locomotion andrespiration. This interaction might be modulated, however, byafferent input from thoracic mechanoreceptors, particularlythe intercostal stretch receptors.     

Do changes in the mechanical properties of articular cartilage promote catabolic destruction of cartilage and osteoarthritis?

Osteoarthritis (OA) is a joint disease characterized by cartilage degeneration, a thickening of subchondral bone, and formation of marginal osteophytes. Previous mechanical characterization of cartilage in our laboratory suggests that energy storage and dissipation is reduced in osteoarthritis as the extent of fibrillation and fissure formation increases. It is not clear whether the loss of energy storage and dissipation characteristics is a result of biochemical and/or biophysical changes that occur to hyaline cartilage in joints. The purpose of this study is to present data, on the strain rate dependence of the elastic and viscous behaviors of cartilage, in order to further characterize changes that occur in the mechanical properties that are associated with OA. We have previously hypothesized that the changes seen in the mechanical properties of cartilage may be due to altered mechanochemical transduction by chondrocytes. Results of incremental tensile stress-strain tests at strain rates between 100%/min and 10,000%/min conducted on OA cartilage indicate that the slope of the elastic stress-strain curve increases with increasing strain rate, unlike the reported behavior of skin and self-assembled collagen fibers. It is suggested that the strain-rate dependence of the elastic stress-strain curve is due to the presence of large quantities of proteoglycans (PGs), which protect articular cartilage by increasing the apparent stiffness. The increased apparent stiffness of articular cartilage at high strain rates may limit the stresses borne and prolong the onset of OA. It is further hypothesized that increased compressive loading of chondrocytes in the intermediate zone of articular cartilage occurs as a result of normal wear to the superficial zone or from excessive impact loading. Once the superficial zone of articular cartilage is worn away, the tension is decreased throughout all cartilage zones leading to increased chondrocyte compressive loading and up-regulation of mechanochemical transduction processes that elaborate catabolic enzymes.     

Mechanical behaviour of in-situ chondrocytes subjected to different loading rates: a finite element study

Experimental findings indicate that in-situ chondrocytes die readily following impact loading, but remain essentially unaffected at low (non-impact) strain rates. This study was aimed at identifying possible causes for cell death in impact loading by quantifying chondrocyte mechanics when cartilage was subjected to a 5% nominal tissue strain at different strain rates. Multi-scale modelling techniques were used to simulate cartilage tissue and the corresponding chondrocytes residing in the tissue. Chondrocytes were modelled by accounting for the cell membrane, pericellular matrix and pericellular capsule. The results suggest that cell deformations, cell fluid pressures and fluid flow velocity through cells are highest at the highest (impact) strain rate, but they do not reach damaging levels. Tangential strain rates of the cell membrane were highest at the highest strain rate and were observed primarily in superficial tissue cells. Since cell death following impact loading occurs primarily in superficial zone cells, we speculate that cell death in impact loading is caused by the high tangential strain rates in the membrane of superficial zone cells causing membrane rupture and loss of cell content and integrity.     

Failure of accentuated vertical body movements to induce cardiac-locomotor coupling.

Cardiac-locomotor coupling (CLC) has been reported during a variety of rhythmic human activities. One reason postulated for such coupling is that axial movements of the viscera during some activities (the "visceral piston") may enhance expulsion of blood from the heart; if so, accentuated vertical movements of the body should provide a powerful stimulus to coupling. To test this hypothesis, we studied 20 subjects hopping and 20 others skipping rope for greater than or equal to 10 min while electrocardiographic and force-platform signals were recorded, from which we derived the subjects' exercise and heart rates. The incidence and intensity of apparent coupling in the test subjects were compared with those of cross-over controls, where the heart rate of each subject was related to the hopping or skipping rate of a matched subject. Ratios consistent with coupling were seen in 10 (50%) hopping subjects under test conditions and in 13 (65%) under control conditions; among skipping subjects, the incidences were 11 (55%) and 10 (50%). In neither group of subjects was the difference in the incidences or the intensities of apparent CLC statistically significant. Our failure to detect CLC while our subjects were hopping or skipping suggests that the visceral piston is unimportant to the CLC phenomenon.     

Applied horizontal force increases impact loading in reduced-gravity running

The chronic exposure of astronauts to microgravity results in structural degradation of their lower limb bones. Currently, no effective exercise countermeasure exists. On Earth, the impact loading that occurs with regular locomotion is associated with the maintenance of bone's structural integrity, but impact loads are rarely experienced in space. Accurately mimicking Earth-like impact loads in a reduced-gravity environment should help to reduce the degradation of bone caused by weightlessness. We previously showed that running with externally applied horizontal forces (AHF) in the anterior direction qualitatively simulates the high-impact loading associated with downhill running on Earth. We hypothesized that running with AHF at simulated reduced gravity would produce impact loads equal to or greater than values experienced during normal running at Earth gravity. With an AHF of 20% of gravity-specific body weight at all gravity levels, impact force peaks increased 74%, average impact loading rates increased 46%, and maximum impact loading rates increased 89% compared to running without any AHF. In contrast, AHF did not substantially affect active force peaks. Duty factor and stride frequency decreased modestly with AHF at all gravity levels. We found that running with an AHF in simulated reduced gravity produced impact loads equal to or greater than those experienced at Earth gravity. An appropriate AHF could easily augment existing partial gravity treadmill running exercise countermeasures used during spaceflight and help prevent musculoskeletal degradation.     

Predator selectivity alters the effect of dispersal on coexistence among apparent competitors

While the majority of studies on dispersal effects on patterns of coexistence among species in a metacommunity have focused on resource competitors, dispersal in systems with predator–prey interactions may provide very different results. Here, we use an analytical model to study the effect of dispersal rates on coexistence of two prey species sharing a predator (apparent competition), when the traits of that predator vary. Specifically, we explore the range in immigration rates where apparent competitors are able to coexist, and how that range changes with predator selectivity and efficiency. We find that if the inferior apparent competitor has a higher probability of being consumed, it will require less immigration to invade and to exclude the superior prey as the predator becomes more opportunistic. However, if the inferior apparent competitor has a lower probability of being consumed (and lower growth rates), higher immigration is required for the inferior prey to invade and exclude the superior prey as the predator becomes more opportunistic. We further find that the largest range of immigration rates where prey coexist occurs when predator selectivity is intermediate (i.e. they do not show much bias towards consuming one species or the other). Increasing predator efficiency generally reduces the immigration rates necessary for the inferior apparent competitor to invade and exclude the superior apparent competitor, but also reduces the range of immigration rates where the two apparent competitors can coexist. However, when the superior apparent competitor has a higher probability of being consumed, increased predator efficiency can increase the range of parameters where the species can coexist. Our results are consistent with some of the variation observed in the effect of dispersal on prey species richness in empirical systems with top predators.     

Long-range correlations between DNA bending sites: relation to the structure and dynamics of nucleosomes

It has been established that the precise positioning of nucleosomes on genomic DNA can be achieved, at least for a minority of them, through sequence-dependent processes. However, to what extent DNA sequences play a role in the positioning of the major part of nucleosomes is still debated. The aim of the present study is to examine to what extent long-range correlations (LRC) are related to the presence of nucleosomes. Using the wavelet transform technique, we perform a comparative analysis of the DNA text and of the corresponding bending profiles generated with curvature tables based on nucleosome positioning data. The exploration of a number of eukaryotic and bacterial genomes through the optics of the so-called "wavelet transform microscope" reveals a characteristic scale of 100-200 bp that separates two regimes of different LRC. Here, we focus on the existence of LRC in the small-scale regime (10-200 bp) which are actually observed in eukaryotic genomes, in contrast to their absence in eubacterial genomes. Analysis of viral DNA genomes shows that, like their host's genomes, eukaryotic viruses present LRC but eubacterial viruses do not. There is one exception for genomes of poxviruses (Vaccinia and Melamoplus sanguinipes) which do not replicate in the cell nucleus and do not exhibit LRC. No small-scale LRC are detected in the genomes of all examined RNA viruses, with the exception of retroviruses. These results together with the observation of LRC between particular sequence motifs known to participate in the formation of nucleosomes (e.g. AA dinucleotides) strongly suggest that the 10-200 bp LRC are a signature of the sequence-dependence of nucleosome positioning. Finally, we discuss possible interpretations of these LRC in terms of the physical mechanisms that might govern the positioning and the dynamics of the nucleosomes along the DNA chain through cooperative processes.     

杭州湾湿地不同演替阶段优势物种光合生理生态特性

采用LI-6400光合作用系统测定了杭州湾滨海湿地不同演替阶段6种优势植物-包括早期植物海三棱藨草(Scirpus mariqueter)和糙叶苔草(Carex scabrifolia)、中期植物芦苇(Phragmites communis)和柽柳(Tamarixchinensis)、后期植物白茅(Imperata cylindrica)和旱柳(Salix matsudana)的光合作用光响应曲线(LRC)和CO2响应曲线(A/Ci),并拟合得出多个光合生理指标.结果表明,6种优势植物LRC净光合速率(Pn)大小顺序为海三棱藨草>糙叶苔草>芦苇>柽柳>白茅>旱柳,且早期植物显著大于后期植物(P<0.05);光补偿点(LCP)、光饱和点(LSP)、最大净光合速率(Amax)、和暗呼吸速率(Rd)变化与Pn相同,也表现出演替早期植物>中期植物>后期植物;而表观量子效率(AQY)则表现出相反趋势.由A/Ci曲线可以发现,演替早期优势植物较后期植物具有更低的CO2羧化效率(CCE)和相对较高的CO2补偿点(CCP).可见群落演替与各阶段优势植物的光合生理特征密切相关.     

水稻种子萌发和苗期ABA敏感性的QTL定位分析

植物激素ABA参与不同的生理过程,尤其是在种子发育和非生物逆境的适应都需要ABA的调控.以水稻珍汕97和旱稻IRAT109为亲本的重组自交系群体为材料,分别调查种子发芽和苗期对ABA的敏感程度.种子发芽阶段以ABA处理下的相对发芽势（Relative germination vigor,RGV）和相对发芽率（Relative germination rate,RGR）为指标,苗期以ABA喷施处理下的卷叶程度（Leaf rolling scores by ABA spaying,LRS）和含ABA水培条件下的卷叶程度（Leaf rolling scores by ABA culturing,LRC）为指标.性状相关分析表明发芽阶段的相对发芽势与苗期卷叶程度呈显著正相关.用复合区间作图法和混合线性模型对ABA敏感性QTL定位和上位性效应分析.两种软件检测到的主效QTL位点大致相同.共检测到5个单位点QTL和6对上位性QTL与发芽阶段的ABA敏感性有关;8个单位点QTL和5对上位性QTL与水稻苗期对ABA的敏感性有关;在苗期,两种ABA处理条件下共检测到两个共同的QTL;仅一个共同的QTL同时控制发芽阶段和苗期对ABA的敏感性.这些研究结果说明,水稻对ABA的敏感性同时受单位点的多基因和上位性基因控制;而且控制种子萌发阶段发芽势和苗期对ABA敏感性的遗传基础有很大的不同.     

The effects of arm crank training on the physiological responses to submaximal wheelchair ergometry

The purpose of this investigation was to examine the cardiovascular and metabolic effects of a 5 wk arm crank (AC) training program on submaximal wheelchair (WC) ergometry in able-bodied women. The 6 subjects in the training group (TG) and 4 in the control group (CG) performed a 10 min WC exercise prior to and following the training period at a power output (PO) that elicited 70% of the pre-training peak oxygen uptake (VO2). Steady state VO2, heart rate (HR), cardiac output (Qc) and stroke volume (Vs) were measured. Resting and post-exercise blood lactate concentrations (LA) were measured, the difference was recorded as net LA. The TG exercised on the AC 3 d.wk-1 at a PO that elicited 85% of each subject's recorded peak HR. Each session consisted of four 4 min exercise bouts preceded by a 2 min warm-up and interspersed with 2 min rest periods. After training, the TG had a significantly (p less than 0.05) lower HR, larger Vs and lower LA in response to the WC exercise. Qc and VO2 were not significantly altered. The results demonstrate that the AC exercise program used in this study produced a physiological training effect which was observed during submaximal WC exercise of an intensity frequently encountered during daily WC ambulation. It appears that short-term, moderate intensity AC training offers an adequate stimulus to reduce the stress imposed by wheelchair locomotion.     

Global patterns of the double mutualism phenomenon

A double mutualism (DM) occurs when two interacting species benefit each other in two different functions, e.g. when an animal species acts both as pollinator and seed disperser of the same plant. Besides the double benefit, a DM also imposes a larger risk to both functions if the performance of one partner declines. We conducted the first global review of DMs involving pollinators and seed dispersers, aiming to: 1) assess their prevalence across ecosystems and biogeographical regions; 2) identify the main plant and animal taxa, and their traits, implicated in DMs; and 3) evaluate the conservation status of double mutualist species. We compiled published and unpublished DM records using specific search terms, noting the species involved, their conservation status and geographic location, as well as the type of study (species vs community‐level) in which the DM was detected. We identified 302 DM cases involving 207 plant and 92 animal species from 16 mainland and 17 island areas. Most records come from tropical regions and islands. Animals included birds (62%), mammals (22%), and reptiles (16%), mostly opportunist species; only 18% were nectarivores. Plants were mainly fleshy‐fruited shrub or tree species (59%) with actinomorphic flowers that were visited by several or many pollinator species (87%). Most (56%) DMs were detected in community‐level studies. DMs are mostly prevalent in ecosystems with limited food resources and mutualist partners, and with high generalization levels. Nearly 30% of the species involved in DMs are threatened according to IUCN criteria, 68% of which are found on islands. The high prevalence of DM on islands paired with the threat status of island species suggest that the loss of a double mutualists and its cascading consequences may have a severe impact on community composition and functioning of fragile island ecosystems.     

Evaluation of the influence of strain rate on Colles' fracture load

Colles' fracture, a transverse fracture of the distal radius bone, is one of the most frequently observed osteoporotic fractures resulting from low energy or traumatic events, associated with low and high strain rates, respectively. Although experimental studies on Colles' fracture were carried out at various loading rates ranging from static to impact loadings, there is no systematic study in the literature that isolates the influence of strain rate on Colles' fracture load. In order to provide a better understanding of fracture risk, the current study combines experimental material property measurements under varying strain rates with computational modeling and presents new information on the effect of strain rate on Colles' fracture. The simulation results showed that Colles' fracture load decreased with increasing strain rate with a steeper change in lower strain rates. Specifically, strain rate values (0.29s(-1)) associated with controlled falling without fracture corresponded to a 3.7% reduction in the fracture load. On the other hand, the reduction in the fracture load was 34% for strain rate of 3.7s(-1) reported in fracture inducing impact cadaver experiments. Further increase in the strain rate up to 18s(-1) led to an additional 22% reduction. The most drastic reduction in fracture load occurs at strain rates corresponding to the transition from controlled to impact falling. These results are particularly important for the improvement of fracture risk assessment in the elderly because they identify a critical range of loading rates (10-50mm/s) that can dramatically increase the risk of Colles' fracture.     

Identification of Quantitative Trait Loci for ABA Sensitivity at Seed Germination and Seedling Stages in Rice

Abscisic acid (ABA) is one of the important plant hormones, which plays a critical role in seed development and adaptation to abiotic stresses. The sensitivity of rice (Oryza sativa L.) to exogenous ABA at seed germination and seedling stages was investigated in the recombinant inbred line (RIL) population derived from a cross between irrigated rice Zhenshan 97 and upland rice IRAT109, using relative germination vigor (RGV), relative germination rate (RGR) and leaf rolling scores of spraying (LRS) or culturing (LRC) with ABA as sensitivity indexes. The phenotypic correlation analysis revealed that only RGV at germination stage was positively correlated to ABA sensitivity at seedling stage. QTL detection using composite interval mapping (CIM) and mixed linear model was conducted to dissect the genetic basis of ABA sensitivity, and the single-locus QTLS detected by both methods are in good agreement with each other. Five single QTLs and six pairs of epistatic QTLs were detected for ABA sensitivity at germination stage. Eight single QTLs and five pairs of epistatic QTLs were detected for ABA sensitivity at seedling stage. Two QTLs were common between LRS and LRC; and one common QTL was detected for RGV, LRS and LRC simultaneously. These results indicated that both single and epistatic loci were involved in the ABA sensitivity in rice, and the genetic basis of ABA sensitivity at seed germination and seedling stage was largely different.     

Effect of locomotor respiratory coupling on respiratory evaporative heat loss in the sheep.

During galloping, many animals display 1:1 coupling of breaths and strides. Locomotor respiratory coupling (LRC) may limit respiratory evaporative heat loss (REHL) by constraining respiratory frequency (f). Five sheep were exercised twice each, according to a five-step protocol: 5 min at the walk, 5 min at the trot (trot1), 10 min at the gallop, 5 min at the trot (trot2), and 5 min at the walk. Rectal temperature (T(re)), stride frequency, f, REHL, and arterial CO(2) tension and pH were measured at each step. Tidal volume (VT) was calculated. LRC was observed only during galloping. The coupling ratio remained at 1:1 while VT increased continuously during galloping, causing REHL to increase from 2.9 +/- 0.2 (SE) W/kg at the end of trot1 to a peak of 5.3 +/- 0.3 W/kg. T(re) rose from 39.0 +/- 0.1 degrees C preexercise to 40.2 +/- 0.2 degrees C at the end of galloping. At the gallop-trot2 transition, VT fell and f rose, despite a continued rise in T(re). Arterial CO(2) tension fell from 36.5 +/- 1.1 Torr preexercise to 31.8 +/- 1.4 Torr by the end of trot1 and then further to 21.5 +/- 1.2 Torr by the end of galloping, resulting in alkalosis. In conclusion, LRC did not prevent increases in REHL in sheep because VT increased. The increased VT caused hypocapnia and presumably elevated the cost of breathing.     

True pedigree errors more frequent than apparent errors for single nucleotide polymorphisms

Single nucleotide polymorphisms (SNPs) are currently being developed for use in disequilibrium analyses. These SNPs consist of two alleles with varying degrees of polymorphism. A natural design for use with SNPs is the 'haplotype relative risk' sampling design in which a father, mother, and child are typed at an SNP locus. Given such a trio of genotypes, we ask: what is the probability that a pedigree error (a change from one allele to the other) at an SNP locus will be detected using only Mendel's laws as a check? We calculate the probability of detecting such errors for a hypothetical SNP locus with varying degrees of polymorphism and for various true error rates. For the sets of allele frequencies considered, we find that the detection rates range between 25 and 30%, the detection rate being lowest when the two alleles have equal frequencies and the highest when one allele has a frequency of 10%. Based on this detection rate, we determine that the true error rate is roughly 3.3-4 times that of the apparent error rate at an SNP locus. The greatest discrepancy between true and apparent error rates occurs when allele frequencies are equal.     

Big plants—do they limit species coexistence?

Aims According to conventional theory, larger plant species are likely to inflict more intense competition on other (smaller) species. We tested a deducible prediction from this: that a larger species should generally be expected to impose greater limits on the number of species that can coexist with it.Methods Species richness was sampled under plant canopies for a selection of woody species ('host' species) that display a wide range of adult sizes (from small shrubs to large trees), growing within natural vegetation of the Interior Douglas-fir zone of southern British Columbia, Canada. These data were compared with species richness levels sampled within randomly placed plots within the host species habitat.Important findings A prominent host species size effect on species richness was detected but only narrowly at the small end of the species size range. Across most (90%) of the increasing size range of host species, the number of species residing under the host canopy showed no significant decrease relative to the number expected by random assembly, based on species richness within randomly defined equivalent areas within the habitat of the host species. This apparent 'null effect', we suggest, is explained not because these larger species have no effect on community assembly. We postulate that larger species are indeed likely to be more effective in causing competitive exclusion of some smaller species (as expected from conventional theory), but that any potential limitation effect of this on resident species richness is offset for two reasons: (i) larger species also generate niche spaces that they cannot exploit under their own canopies and so have minimal impact (as competitors) on smaller species that can occupy these niches and (ii) certain other small species—despite small size—have effective competitive abilities under the severe competition that occurs within host neighbourhoods of larger species. These and other recent studies call for re-evaluation of traditional views on the role of plant size in affecting competitive ability and community assembly.     

Sex-ratio adjustment when relatives interact: a test of constraints on adaptation

Studies of sex allocation offer excellent opportunities for examining the constraints and limits on adaptation. A major topic of debate within this field concerns the extent to which the ability of individuals to adaptively manipulate their offspring sex ratio is determined by constraints such as the method of sex determination. We address this problem by comparing the extent of sex-ratio adjustment across taxa with different methods of sex determination, under the common selective scenario of interactions between relatives. These interactions comprise the following: local resource competition (LRC), local mate competition (LMC), and local resource enhancement (LRE). We found that: (1) species with supposedly constraining methods of sex determination showed consistent sex-ratio adjustment in the predicted direction; (2) vertebrates with chromosomal sex determination (CSD) showed less adjustment then haplodiploid invertebrates; (3) invertebrates with possibly constraining sex-determination mechanisms (CSD and pseudo-arrhenotoky) did not show less adjustment then haplodiploid invertebrates; (4) greater sex-ratio adjustment was seen in response to LRC and LMC than LRE; (5) greater sex-ratio adjustment was seen in response to interactions between relatives (LRC, LMC, and LRE) compared to responses to other environmental factors. Our results also illustrate the problem that sex-determination mechanism and selective pressure are confounded across taxa because vertebrates with CSD are influenced primarily by LRE whereas invertebrates are influenced by LRC and LMC. Overall, our analyses suggest that sex-allocation theory needs to consider simultaneously the influence of variable selection pressures and variable constraints when applying general theory to specific cases.     

The incidence and diversity of Wolbachia in gallwasps (Hymenoptera; Cynipidae) on oak

Wolbachia bacteria infect approximately 20% of all insect species, and cause a range of alterations to host reproduction, including imposition of thelytoky. The incidence and phenotypic impact of Wolbachia remains to be established in many insect taxa, and considerable research effort is currently focused on its association with particular reproductive modes and the relative importance of the various pathways via which infection occurs. Gallwasps represent an attractive system for addressing these issues for two reasons. First, they show a diversity of reproductive modes (including arrhenotoky, thelytoky and cyclical parthenogenesis) in which the impact of Wolbachia infection can be examined. Second, they occupy two intimately linked trophic niches (gall-inducers and inquilines) between which there is potential for the horizontal exchange of Wolbachia infection. In the arrhenotokous gallwasp lineages screened to date (the herb-galling 'Aylacini' and the rose-galling Diplolepidini), Wolbachia infection always induces thelytoky. The impact of Wolbachia in other arrhenotokous clades, and in the cyclically parthenogenetic clades remains unknown. Here we use polymerase chain reaction (PCR) screening and sequence data for two Wolbachia genes (wsp and ftsZ) to examine the prevalence and incidence of Wolbachia infection in 64 species (a total of 609 individuals) in two further tribes: the arrhenotokous inquilines (tribe Synergini), and the cyclically parthenogenetic oak gallwasps (tribe Cynipini). We ask: (i) whether Wolbachia infection has any apparent impact on host reproduction in the two tribes and (ii) whether there is any correlation between Wolbachia infection and the apparent lack of an arrhenotokous generation in many oak gallwasp life cycles. We show: (i) that Wolbachia infection is rare in the Cynipini. Infected species show no deviation from cyclical parthenogenesis, and infection is no more common in species known only from a thelytokous generation; (ii) that there is a higher incidence of infection within the arrhenotokous inquilines, and generally in gallwasp tribes without cyclical parthenogensis; (iii) all Wolbachia-positive inquiline species are known to possess males, implying either that Wolbachia infection does not result in loss of sex in this tribe or, more probably, that (as for some rose gallwasps) Wolbachia infection leads to loss of sex in specific populations; and (iv) although we find some inquilines and gall inducers to be infected with Wolbachia having the same wsp sequence, these hosts are not members of the same gall communities, arguing against frequent horizontal transmission between these two trophic groups. We suggest that exchange may be mediated by the generalist parasitoids common in oak galls.     

Floor anchorage load and safety space for adult wheelchair users during a crash

Over the last 20 years in France as well as generally in Europe and also in North America, the integration of disabled people into society has become a more and more pressing issue. Work is in progress to increase the safety level of wheelchair occupants up to becoming equivalent to that of able-bodied road vehicles occupants. Present world practices are that wheelchair occupants who are unable to transfer to a vehicle seat for transportation remain in their wheelchair, which is secured to the vehicle floor using tie downs, and the occupant is restrained in the wheelchair. The aim of this paper was to perform a series of crash tests to make sure that the current standards are suitable to identify possible failures in safety in the restraint systems intended for wheelchair users. To date, there is no specific regulatory criterion related to floor anchorage for wheelchairs in vehicles during a crash. At a 48 km/h crash test, loads reached 30 kN on rear anchorage, and 13 kN on shoulder belt. Also the kinematics analysis of the wheelchair user during impact pointed out dimension of his safety space.