Coupling of posture and gait: mode locking and parametric excitation.

We investigate the temporal coordination of human gait and posture and infer the nature of their coupling. Participants viewed a sinusoidally oscillating visual display which induced medial-lateral postural sway during treadmill walking, while display frequency was varied (0.075-1.025 Hz). First, postural responses exhibited the usual low-pass characteristic but with an additional resonance peak near the preferred stride frequency, although shifted downward by 0.12 Hz; this provides evidence of a coupling from gait to posture. Second, the step cycle adapted to mode lock with the visual driver and postural sway, as well as displaying instances of intermittency (slipping in and out of phase) and quasiperiodicity (phase wandering); this provides evidence of a coupling from posture to gait. We observed a spectrum of integer mode locks, including a large 1:1 trapping region about the stride frequency and superharmonic entrainment (stride frequency > driver frequency) at lower driver frequencies. A coupled-oscillator model that incorporates a novel parametric coupling from posture to the gait "stiffness" term reproduces these features of the data, including the resonance peak shift. Biological coordination patterns may thus emerge naturally as properties of a system of appropriately coupled oscillators.

Optic flow is used to control human walking.

How is human locomotion visually controlled? Fifty years ago, it was proposed that we steer to a goal using optic flow, the pattern of motion at the eye that specifies the direction of locomotion. However, we might also simply walk in the perceived direction of a goal. These two hypotheses normally predict the same behavior, but we tested them in an immersive virtual environment by displacing the optic flow from the direction of walking, violating the laws of optics. We found that people walked in the visual direction of a lone target, but increasingly relied on optic flow as it was added to the display. The visual control law for steering toward a goal is a linear combination of these two variables weighted by the magnitude of flow, thereby allowing humans to have robust locomotor control under varying environmental conditions.

Haptic interaction with virtual objects. Spatial perception and motor control.

This paper considers interaction of the human arm with "virtual" objects simulated mechanically by a planar robot. Haptic perception of spatial properties of objects is distorted. It is reasonable to expect that it may be distorted in a geometrically consistent way. Three experiments were performed to quantify perceptual distortion of length, angle and orientation. We found that spatial perception is geometrically inconsistent across these perceptual tasks. Given that spatial perception is distorted, it is plausible that motor behavior may be distorted in a way consistent with perceptual distortion. In a fourth experiment, subjects were asked to draw circles. The results were geometrically inconsistent with those of the length perception experiment. Interestingly, although the results were inconsistent (statistically different), this difference was not strong (the relative distortion between the observed distributions was small). Some computational implications of this research for haptic perception and motor planning are discussed.

The role of central and peripheral vision in postural control during walking.

Three hypotheses have been proposed for the roles of central and peripheral vision in the perception and control of self-motion: (1) peripheral dominance, (2) retinal invariance, and (3) differential sensitivity to radial flow. We investigated postural responses to optic flow patterns presented at different retinal eccentricities during walking in two experiments. Oscillating displays of radial flow (0 degree driver direction), lamellar flow (90 degrees), and intermediate flow (30 degrees, 45 degrees) patterns were presented at retinal eccentricities of 0 degree, 30 degrees, 45 degrees, 60 degrees, or 90 degrees to participants walking on a treadmill, while compensatory body sway was measured. In general, postural responses were directionally specific, of comparable amplitude, and strongly coupled to the display for all flow patterns at all retinal eccentricities. One intermediate flow pattern (45 degrees) yielded a bias in sway direction that was consistent with triangulation errors in locating the focus of expansion from visible flow vectors. The results demonstrate functionally specific postural responses of both central and peripheral vision, contrary to the peripheral dominance and differential sensitivity hypotheses, but consistent with retinal invariance. This finding emphasizes the importance of optic flow structure for postural control regardless of the retinal locus of stimulation.

Phenotypic diversity of enterotoxigenic Escherichia coli strains from a community-based study of pediatric diarrhea in periurban Egypt.

No past studies of diarrhea in children of the Middle East have examined in detail the phenotypes of enterotoxigenic Escherichia coli (ETEC) strains, which are important pathogens in this setting. During a prospective study conducted from November 1993 to September 1995 with 242 children under 3 years of age with diarrhea living near Alexandria, Egypt, 125 episodes of diarrhea were positive for ETEC. ETEC strains were available for 98 of these episodes, from which 100 ETEC strains were selected and characterized on the basis of enterotoxins, colonization factors (CFs), and O:H serotypes. Of these representative isolates, 57 produced heat-stable toxin (ST) only, 34 produced heat-labile toxin (LT) only, and 9 produced both LT and ST. Twenty-three ETEC strains expressed a CF, with the specific factors being CF antigen IV (CFA/IV; 10 of 23; 43%), CFA/II (5 of 23; 22%), CFA/I (3 of 23; 13%), PCFO166 (3 of 23; 13%), and CS7 (2 of 23; 9%). No ETEC strains appeared to express CFA/III, CS17, or PCFO159. Among the 100 ETEC strains, 47 O groups and 20 H groups were represented, with 59 O:H serotypes. The most common O serogroups were O159 (13 strains) and O43 (10 strains). O148 and O21 were each detected in five individual strains, O7 and O56 were each detected in four individual strains, O73, O20, O86, and O114 were each detected in three individual strains, and O23, O78, O91, O103, O128, and O132 were each detected in two individual strains. The most common H serogroups were H4 (16 strains), 12 of which were of serogroup O159; H2 (9 strains), all of which were O43; H18 (6 strains); H30 (6 strains); and H28 (5 strains); strains of the last three H serogroups were all O148. Cumulatively, our results suggest a high degree of clonal diversity of disease-associated ETEC strains in this region. As a low percentage of these strains expressed a CF, it remains possible that other adhesins for which we either did not assay or that are as yet undiscovered are prevalent in this region. Our findings point out some potential barriers to effective immunization against ETEC diarrhea in this population and emphasize the need to identify additional protective antigens commonly expressed by ETEC for inclusion in future vaccine candidates.

Identification of Shigella flexneri subserotype 1c in rural Egypt.

In a population-based study of diarrhea in rural, northern Egypt, 60 Shigella flexneri strains were identified, of which 10 could not be definitively serotyped. Serological analysis with commercial reagents suggested that they were serotype 1, but the strains failed to react with subserotype 1a- or 1b-specific antibodies. All 10 strains reacted with MASF 1c, a monoclonal antibody specific for a provisional S. flexneri subserotype, 1c, first identified in Bangladesh and not previously detected outside of that region. Our results show that S. flexneri subserotype 1c is not unique to Bangladesh and that the inability to detect it may reflect both the limited use of suitable screening methods and the rarity of this subserotype.

Epidemiology of enterotoxigenic Escherichia coli diarrhea in a pediatric cohort in a periurban area of lower Egypt.

Enterotoxigenic Escherichia coli (ETEC) are diverse pathogens that express heat-labile (LT) and/or heat-stable (ST) enterotoxins, yet little is known about whether epidemiologic patterns of pediatric ETEC diarrhea vary by the expressed ETEC toxin phenotype. In total, 242 Egyptian children aged <3 years were prospectively followed in 1993-1995. ETEC episodes were detected during twice-weekly home visits, and asymptomatic ETEC excretion was identified from monthly cross-sectional surveys. ETEC episodes were 0.6 per child-year. ST-only ETEC was 2.6 times (P<.001) more common in warmer than cooler months, while LT-only ETEC showed no seasonal variation. Ownership of a household sanitary latrine, but not breast-feeding, was associated with a lower risk of both enterotoxin phenotypes. Coexpression of a colonization factor by LT- or ST-only ETEC strengthened the association with diarrhea. These findings indicate that the epidemiologic patterns of LT-only and ST-only ETEC are not identical and that disease interventions should include improved household sanitation.

Motion parallax is used to control postural sway during walking.

Three experiments tested the hypothesis that postural sway during locomotion is visually regulated by motion parallax as well as optical expansion. Oscillating displays of three-dimensional scenes were presented to participants walking on a treadmill, while postural sway was recorded. Displays simulated: (a) a cloud, in which parallax and expansion are congruent, (b) a hallway, (c) the side walls of the hallway, (d) a ground surface, (e) a wall, (f) the wall with a central hole, (g) a hall farther from the observer, and (h) a wall farther from the observer. In contrast to previous results with a hallway, responses with the cloud were isotropic and directionally specific. The other displays demonstrated that motion parallax was more effective than simple horizontal flow in eliciting lateral sway. These results are consistent with the hypothesis that adaptive control of sway during walking is based on congruent expansion and parallax in natural environments.

Visual control of posture during walking: functional specificity.

Three experiments examined the functional specificity of visually controlled posture during locomotion by presenting large-screen displays to participants walking on a treadmill. Displays simulated locomotion down a stationary hallway, a hallway that traveled with the observer, or a frontal wall that traveled with the observer. A superimposed oscillation specified postural sway in 6 possible directions. With the wall, sway amplitude was isotropic and directionally specific in all conditions. However, with the hallways, sway was anisotropic (lateral > anterior-posterior [AP]), and diagonal responses were flattened into the lateral plane. When the treadmill was turned 90 degrees to the hallway, both the anisotropy and flattening were reversed (AP > lateral), indicating that they are determined by the visual structure of the scene. The results can be explained by postural control laws based on both optical expansion and motion parallax, yielding biases in planar environments that truncate parallax.

Field trial of inactivated oral cholera vaccines in Bangladesh: results from 5 years of follow-up.

To determine the protective efficacy (PE) of three doses of oral B subunit-killed whole cell (BS-WC) or killed whole cell-only (WC) vaccines against cholera, a clinical trial was conducted among 62285 children over 2 years and adult women in rural Bangladesh. During 5 years of follow-up, there were 144 cases of cholera in the BS-WC group (PE = 49%; P < 0.001), 150 in the WC group (PE = 47%; P < 0.001), and 283 in the K12 group. Protection by each vaccine was evident only during the first three years of follow-up; long-term protection of young children was observed only against classical but not El Tor cholera; 3-year protection against both cholera biotypes occurred among older persons, but at a higher level against classical cholera.

Infant bouncing: the assembly and tuning of action systems.

We outline a theory of infant skill acquisition characterized by an assembly phase, during which a task-specific, low-dimensional action pattern emerges from spontaneous movement in the context of task constraints, and a tuning phase, during which adjustment of the system parameters yields a more energetically efficient and more stable movement. 8 infants were observed longitudinally when bouncing while supported by a harness attached to a spring. We found an initial assembly phase in which kicking was irregular and variable in period, and a tuning phase with more periodic kicking, followed by the sudden appearance of long bouts of sustained bouncing. This "peak" behavior was characterized by oscillation at the resonant frequency of the mass-spring system, an increase in amplitude, and a decrease in period variability. The data are consistent with a forced mass-spring operating at resonance.

Prevalence of Plesiomonas shigelloides among diarrhoeal patients in Bangladesh.

The incidence of Plesiomonas shigelloides among diarrhoeal patients attending the Dhaka Treatment Centre of the International Centre for Diarrhoeal Disease Research, Bangladesh (ICDDR, B) from January through December 1987, has been reported. Using bile peptone broth (pH 8.8) as an enrichment medium and Salmonella-Shigella agar to isolate the organism, P. shigelloides was isolated from 838 (6.4%) of 13,142 patients, 523 (4.0%) of whom had no other pathogen isolated. The percentage of isolation was higher from stool (9.2%) than from rectal swab (5.9%) specimens (P < 0.005). The incidence of P. shigelloides was higher among male (64.0%) than female (36.0%) patients (P < 0.005). Isolation was highest among children less than five years. P. shigelloides was isolated most often in March (11.0%) and September (7.7%), indicating two seasonal peaks of incidence before and after the monsoons. All the strains were uniformly sensitive to chloramphenicol, furazolidon, gentamicin and trimethoprim-sulfamethoxazole. Sensitivity of the organism to ampicillin and tetracycline was 27.0% and 89.0%, respectively. This indicates that P. shigelloides may be an important agent of diarrhoea in our patient population.

DNA probe analysis of diarrhoeagenic Escherichia coli: detection of EAF-positive isolates of traditional enteropathogenic E. coli serotypes among Bangladeshi paediatric diarrhoea patients.

Escherichia coli isolates from all surveillance patients less than or equal to 20 months of age seen for diarrhoea at the Dhaka Clinical Treatment Facility of the International Centre for Diarrhoeal Disease Research, Bangladesh between March 1 and August 31, 1988, were collected and hybridized with DNA probes to assess the potential importance of diarrhoeagenic E. coli among paediatric patients in Bangladesh. Of 396 patients evaluated, 18% were infected with enteropathogenic E. coli (EPEC) adherence factor (EAF)-positive E. coli, 23% were infected with enterotoxigenic E. coli (ETEC), 9% were infected with Shiga-like toxin-positive E. coli, and 13% were infected with diffuse adhesiveness-positive E. coli. None were infected with enteroinvasive E. coli. Ten percent of patients were colonized with more than one type of potential diarrhoeagenic E. coli. The majority of EAF-positive isolates were of traditional EPEC O:H serotypes. Although this was not a case-control study, the large number of EPEC and ETEC, which are recognized enteric pathogens, suggests these organisms are important causes of diarrhoeal diseases in this pediatric population.

Yersinia enterocolitica isolated from two cohorts of young children in Santiago, Chile: incidence of and lack of correlation between illness and proposed virulence factors.

Yersinia enterocolitica was isolated from children in two cohorts in Santiago, Chile. In a cohort containing a cross section of children aged 0 to 4 years, Y. enterocolitica was isolated from stool samples of 1.1% of children with diarrhea and 0.2% of age-matched control children. In a subgroup of this cohort from which weekly stool samples were obtained from all children irrespective of clinical status, 6% of children had asymptomatic Yersinia infections. In a birth cohort (with a greater representation of children less than 1 year of age and a significantly higher rate of diarrhea), Y. enterocolitica was isolated from 1.9% of children with diarrhea and 0.6% of controls (P = 0.05). Biogroup 1A strains (which lacked traditional phenotypic and molecular markers for pathogenicity) were isolated from seven children with diarrhea but from no control children in the birth cohort (P = 0.02). All other isolates, including all isolates from asymptomatic children, were "pathogenic" strains in biogroup 4, serogroup O3; no association between these isolates and occurrence of disease was found. Y. enterocolitica is found among young children in Santiago, with asymptomatic infections not uncommon occurrences. However, questions about the association between previously described virulence factors and diarrheal illness remain.

Biotype as determinant of natural immunising effect of cholera.

To test the hypothesis that clinical Vibrio cholerae O1 infections protect against recurrent cholera, treated cholera episodes in a rural Bangladesh population of 188,153 people who were followed between 1985 and 1988 were analysed. Of the 2214 people with initial episodes of cholera, 7 had a second episode. The incidence of cholera was 61% lower in subjects who had had an earlier episode than in those without such an episode. Whereas initial episodes of classical cholera were associated with complete protection against subsequent cholera, initial episodes of El Tor cholera were associated with negligible protection.

Steady-state and perturbed rhythmical movements: a dynamical analysis.

This study examined rhythmic finger movements in the steady state and when momentarily perturbed in order to derive their qualitative dynamical properties. Movement frequency, amplitude, and peak velocity were stable under perturbation, signaling the presence of an attractor, and the topological dimensionality of that attractor was approximately equal to one. The strength of the attractor was constant with increasing movement frequency, and the Fourier spectra of the steady-state trials showed an alternating harmonic pattern. These results are consistent with a previously derived nonlinear oscillator model. However, the oscillation was phase advanced by perturbation overall, and a consistent phase-dependent, phase-shift pattern occurred, which is inconsistent with the model. The overall phase advance also shows that any central pattern generator responsible for generating the rhythm must be nontrivially modulated by the limb being controlled.

Enterotoxin and cytotoxin production by enteroinvasive Escherichia coli.

It has long been suspected that besides their ability to invade enterocytes, enteroinvasive Escherichia coli (EIEC) strains have the ability to elaborate an enterotoxin. We tested 35 EIEC strains for cytotoxins and 9 (1 per serogroup) for enterotoxins. All 35 strains exhibited low levels of Vero cell cytotoxins that are immunologically and genetically distinct from Shiga-like toxin I or II of enterohemorrhagic E. coli. Sterile supernatants and cell lysates of two EIEC strains were tested in rabbit ileal loops, and both stimulated moderate fluid accumulation (circa 0.5 ml/cm) without tissue damage; secretory activity was confirmed in Ussing chambers, where these two strains and the seven others tested significantly increased short circuit current without altering tissue conductance. Curing the 140-MDa invasiveness plasmid from an EIEC strain did not diminish enterotoxin production. Culture in minimal Fe2+ medium is necessary to detect expression of the enterotoxin which is circa 68 to 80 kDa in size and is distinct from the EIEC cytotoxin.

Trimethoprim resistance gene in Shigella dysenteriae 1 isolates obtained from widely scattered locations of Asia.

Trimethoprim-resistance genes of Shigella dysenteriae 1 strains, isolated from a different location of six different countries of Asia over a 5-year period were characterized by using three different dihydrofolate reductase (DHFR) gene probes. The trimethoprim-resistant (TMPR) strains hybridized only with the type I DHFR gene probe by colony hybridization. None of the strains hybridized with types II and III DHFR gene probes. Southern blot experiments using plasmid DNA extracted from these resistant strains indicated that the type I DHFR genes were either on a 20 MDa plasmid or might be located on the chromosome. None of the other plasmids present in S. dysenteriae 1 strains hybridized with the probe. This indicates that the TMP resistance in these S. dysenteriae 1 strains are mediated by type I DHFR enzyme, and there may be transposition of this type I DHFR gene occurs between the 20 MDa plasmid and the chromosome in this serotype of shigella.

Clostridium perfringens type C in bloody and watery diarrhea in Bangladesh.

A two-week age-matched case-control survey to detect Clostridium perfringens type C-associated illness was conducted among diarrheal patients hospitalized following the consumption of high protein meals on the occasion of an annual religious festival. From the stool of 33 patients screened, C. perfringens type C was isolated from one patient with bloody diarrhea and from two patients with watery diarrhea. In Bangladesh, C. perfringens type C has been isolated in both bloody and watery diarrhoeal illness and is not necessarily associated with necrotizing enteritis (pigbel).