Wave-slope soaring of the brown pelican.

BACKGROUND: From the laboratory at Scripps Institution of Oceanography, it is common to see the brown pelican (Pelecanus occidentalis) traveling along the crests of ocean waves just offshore of the surf-zone. When flying in this manner, the birds can travel long distances without flapping, centimeters above the ocean's surface. Here we derive a theoretical framework for assessing the energetic savings related to this behavior, 'wave-slope soaring,' in which an organism in flight takes advantage of localized updrafts caused by traveling ocean surface gravity waves. METHODS: The energy cost of steady, constant altitude flight in and out of ground effect are analyzed as controls. Potential flow theory is used to quantify the ocean wave-induced wind associated with near-shoaling, weakly nonlinear, shallow water ocean surface gravity waves moving through an atmosphere initially at rest. Using perturbation theory and the Green's function for Laplace's equation in 2D with Dirichlet boundary conditions, we obtain integrals for the horizontal and vertical components of the wave-induced wind in a frame of reference moving with the wave. Wave-slope soaring flight is then analyzed using an energetics-based approach for waves under a range of ocean conditions and the body plan of P. occidentalis. RESULTS: For ground effect flight, we calculate a ∼15 - 25% reduction in cost of transport as compared with steady, level flight out of ground effect. When wave-slope soaring is employed at flight heights ∼2m in typical ocean conditions (2m wave height, 15s period), we calculate 60-70% reduction in cost of transport as compared with flight in ground effect. A relatively small increase in swell amplitude or decrease in flight height allows up to 100% of the cost of transport to be offset by wave-slope soaring behavior. CONCLUSIONS: The theoretical development presented here suggests there are energy savings associated with wave-slope soaring. Individual brown pelicans may significantly decrease their cost of transport utilizing this mode of flight under typical ocean conditions. Thus wave-slope soaring may provide fitness benefit to these highly mobile organisms that depend on patchy prey distribution over large home ranges.

Visualising Silicon in Plants: Histochemistry, Silica Sculptures and Elemental Imaging.

Silicon is a non-essential element for plants and is available in biota as silicic acid. Its presence has been associated with a general improvement of plant vigour and response to exogenous stresses. Plants accumulate silicon in their tissues as amorphous silica and cell walls are preferential sites. While several papers have been published on the mitigatory effects that silicon has on plants under stress, there has been less research on imaging silicon in plant tissues. Imaging offers important complementary results to molecular data, since it provides spatial information. Herein, the focus is on histochemistry coupled to optical microscopy, fluorescence and scanning electron microscopy of microwave acid extracted plant silica, techniques based on particle-induced X-ray emission, X-ray fluorescence spectrometry and mass spectrometry imaging (NanoSIMS). Sample preparation procedures will not be discussed in detail, as several reviews have already treated this subject extensively. We focus instead on the information that each technique provides by offering, for each imaging approach, examples from both silicifiers (giant horsetail and rice) and non-accumulators (Cannabis sativa L.).

The use of columns of the zeolite clinoptilolite in the remediation of aqueous nuclear waste streams.

Mud Hills clinoptilolite has been used in an effluent treatment plant (SIXEP) at the Sellafield nuclear reprocessing site. This material has been used to remove 134/137Cs and 90Sr successfully from effluents for 3 decades. Samples of the zeolite have been tested in column experiments to determine their ability to remove radioactive Cs+ and Sr2+ ions under increasing concentrations of competing ions, Ca2+, Mg2+, Na+ and K+. These ions caused increased elution of Cs+ and Sr2+. Ca2+, Mg2+ and K+ were more effective competitors than Na+. For Na+, it was found that if concentration was reduced, then column performance recovered rapidly.

Is callose required for silicification in plants?

The cell wall polymer callose catalyses the formation of silica in vitro and is heavily implicated in biological silicification in Equisetum (horsetail) and Arabidopsis (thale cress) in vivo Callose, a ß-1,3-glucan, is an ideal partner for silicification, because its amorphous structure and ephemeral nature provide suitable microenvironments to support the condensation of silicic acid into silica. Herein, using scanning electron microscopy, immunohistochemistry and fluorescence, we provide further evidence of the cooperative nature of callose and silica in biological silicification in rice, an important crop plant and known silica accumulator. These new data along with recently published research enable us to propose a model to describe the intracellular events that together determine callose-driven biological silicification.

Rough and tough. How does silicic acid protect horsetail from fungal infection?

Horsetail (Equisetum arvense) plants grew healthily for 10 weeks under both Si-deficient and Si-replete conditions. After 10 weeks, plants grown under Si-deficient conditions succumbed to fungal infection. We have used NanoSIMS and fluorescence microscopy to investigate silica deposition in the tissues of these plants. Horsetail grown under Si-deficient conditions did not deposit identifiable amounts of silica in their tissues. Plants grown under Si-replete conditions accumulated silica throughout their tissues and especially in the epidermis of the outer side of the leaf and the furrow region of the stem where it was continuous and often, as a double layer suggestive of a barrier function. We have previously shown, both in vivo (in horsetail and thale cress) and in vitro (using an undersaturated solution of Si(OH)4), that callose is a "catalyst" of plant silica deposition. Here we support this finding by comparing the deposition of silica to that of callose and by showing that they are co-localized. We propose the existence of a synergistic mechanical protection by callose and silica against pathogens in horsetail, whereby the induction of callose synthesis and deposition is the first, biochemical line of defence and callose-induced precipitation of silica is the second, adventitious mechanical barrier.

A database of lumbar spinal mechanical behavior for validation of spinal analytical models.

Data from two experimental studies with eight specimens each of spinal motion segments and/or intervertebral discs are presented in a form that can be used for comparison with finite element model predictions. The data include the effect of compressive preload (0, 250 and 500N) with quasistatic cyclic loading (0.0115Hz) and the effect of loading frequency (1, 0.1, 0.01 and 0.001Hz) with a physiological compressive preload (mean 642N). Specimens were tested with displacements in each of six degrees of freedom (three translations and three rotations) about defined anatomical axes. The three forces and three moments in the corresponding axis system were recorded during each test. Linearized stiffness matrices were calculated that could be used in multi-segmental biomechanical models of the spine and these matrices were analyzed to determine whether off-diagonal terms and symmetry assumptions should be included. These databases of lumbar spinal mechanical behavior under physiological conditions quantify behaviors that should be present in finite element model simulations. The addition of more specimens to identify sources of variability associated with physical dimensions, degeneration, and other variables would be beneficial. Supplementary data provide the recorded data and Matlab® codes for reading files. Linearized stiffness matrices derived from the tests at different preloads revealed few significant unexpected off-diagonal terms and little evidence of significant matrix asymmetry.

Adolescent idiopathic scoliosis.

Adolescent idiopathic scoliosis (AIS) is the most common form of structural spinal deformities that have a radiological lateral Cobb angle - a measure of spinal curvature - of ≥10(°). AIS affects between 1% and 4% of adolescents in the early stages of puberty and is more common in young women than in young men. The condition occurs in otherwise healthy individuals and currently has no recognizable cause. In the past few decades, considerable progress has been made towards understanding the clinical patterns and the three-dimensional pathoanatomy of AIS. Advances in biomechanics and technology and their clinical application, supported by limited evidence-based research, have led to improvements in the safety and outcomes of surgical and non-surgical treatments. However, the definite aetiology and aetiopathogenetic mechanisms that underlie AIS are still unclear. Thus, at present, both the prevention of AIS and the treatment of its direct underlying cause are not possible.

Metabolic Effects of Angulation, Compression, and Reduced Mobility on Annulus Fibrosis in a Model of Altered Mechanical Environment in Scoliosis.

STUDY DESIGN: Comparison of disc tissue from rat tails in 6 groups with different mechanical conditions imposed. OBJECTIVES: To identify disc annulus changes associated with the supposed altered biomechanical environment in a spine with scoliosis deformity using an immature rat model that produces disc narrowing and wedging. BACKGROUND: Intervertebral discs become wedged and narrowed in a scoliosis curve, probably partly because of an altered biomechanical environment. METHODS: We subjected tail discs of 5-week-old immature Sprague-Dawley rats to an altered mechanical environment using an external apparatus applying permutations of loading and deformity for 5 weeks. Together with a sham and a control group, we studied 4 groups of rats: A) 15° angulation, B) angulation with 0.1 MPa compression, C) 0.1 MPa compression, and R) reduced mobility. We measured disc height changes and matrix composition (water, deoxyribonucleic acid, glycosaminoglycan, and hyaluronic acid content) after 5 weeks, and proline and sulphate incorporation and messenger ribonucleic acid expression at 5 days and 5 weeks. RESULTS: After 5 weeks, disc space was significantly narrowed relative to internal controls in all 4 intervention groups. Water content and cellularity (deoxyribonucleic acid content) were not different at interventional levels relative to internal controls and not different between the concave and convex sides of the angulated discs. There was increased glycosaminoglycan content in compressed tissue (in Groups B and C), as expected, and compression resulted in a decrease in hyaluronic acid size. We observed slightly increased incorporation of tritiated proline into the concave side of angulated discs and compressed discs. Asymmetries of gene expression in Groups A and B and some group-wise differences did not identify consistent patterns associating the discs' responses to mechanical alterations. CONCLUSIONS: Intervertebral discs in this model underwent substantial narrowing after 5 weeks, with minimal alteration in tissue composition and minimal evidence of metabolic changes.

Contributions of Remodeling and Asymmetrical Growth to Vertebral Wedging in a Scoliosis Model.

STUDY DESIGN: We performed a laboratory study of rats of 3 different ages with imposed angulation and compressive loading to caudal vertebrae to determine causes of vertebral wedging. OBJECTIVES: The purpose was to determine the percentage of total vertebral wedging that was caused by asymmetric growth, vertebral body, and epiphyseal wedging. Approval from the Institutional Animal Care and Use Committee, the University of Vermont, was obtained for the live animal procedures used in this study. BACKGROUND SUMMARY: Vertebral wedging from asymmetrical growth (Hueter-Volkmann law) is reported to cause vertebral wedging in scoliosis with little attention to the possible contribution of bony remodeling (Wolff's law). METHODS: In our study, an external fixator imposed a 30° lateral curvature and compression of 0.1 megapascal (MPa) in 5- and 14-week-old animals (Groups 1 and 2) and 0.2 MPa in 14- and 32-week-old animals (groups 3 and 4). Total vertebral wedging was measured from micro CT scans. Wedging due to asymmetrical growth and epiphyseal remodeling was calculated from fluorescent labels and the difference was attributed to vertebral body wedging. RESULTS: Total vertebral wedging averaged 18°, 6°, 10° and 5° in Groups 1, 2, 3, and 4, respectively. Metaphyseal asymmetrical growth averaged 8°, 1°, 4°, 0° (44%, 17%, 40% and 0% of total). Epiphyseal wedging averaged 9°, 0°, 3°, and -1°. The difference (vertebral body) averaged 1°, 5°, 3°, and 7° (6%, 83%, 30% and 140% of total). The growth of the loaded vertebrae as a percentage of control vertebrae was 56%, 39% and 25% in Groups 1, 2 and 3; negligible in Group 4. Vertebral body cortical remodeling, with increased thickness and increased curvature on the concave side was evident in young animals and 0.2 MPa loaded older animals. CONCLUSIONS: We conclude that asymmetrical growth was the largest contributor to vertebral wedging in young animals; vertebral body remodeling was the largest contributor in older animals. If, conversely, vertebral wedging can be corrected by appropriate loading in young and old animals, it has important implications for the nonfusion treatment of scoliosis.

Intervertebral disc changes with angulation, compression and reduced mobility simulating altered mechanical environment in scoliosis.

PURPOSE: The intervertebral discs become wedged and narrowed in scoliosis, and this may result from altered biomechanical environment. The effects of four permutations of disc compression, angulation and reduced mobility were studied to identify possible causes of progressive disc deformity in scoliosis. The purpose of this study was to document morphological and biomechanical changes in four different models of altered mechanical environment in intervertebral discs of growing rats and in a sham and control groups. METHODS: External rings were attached by percutaneous pins transfixing adjacent caudal vertebrae of 5-week-old Sprague-Dawley rats. Four experimental Groups of animals underwent permutations of the imposed mechanical conditions (A) 15° disc angulation, (B) angulation with 0.1 MPa compression, (C) 0.1 MPa compression and (R) reduced mobility (N = 20 per group), and they were compared with a sham group (N = 12) and control group (N = 8) (total of 6 groups of animals). The altered mechanical conditions were applied for 5 weeks. Intervertebral disc space was measured from micro-CT images at weeks 1 and 5. Post euthanasia, lateral bending stiffness of experimental and within-animal control discs was measured in a mechanical testing jig and collagen crimp was measured from histological sections. RESULTS: After 5 weeks, micro-CT images showed disc space loss averaging 35, 53, 56 and 35% of the adjacent disc values in the four intervention groups. Lateral bending stiffness was 4.2 times that of within-animal controls in Group B and 2.3 times in Group R. The minimum stiffness occurred at an angle close to the in vivo value, indicating that angulated discs had adapted to the imposed deformity, this is also supported by measurements of collagen crimping at concave and convex sides of the disc annuli. CONCLUSION: Loss of disc space was present in all of the instrumented discs. Thus, reduced mobility, that was common to all interventions, may be a major source of the observed disc changes and may be a factor in disc deformity in scoliosis. Clinically, it is possible that rigid bracing for control of scoliosis progression may have secondary harmful effects by reducing spinal mobility.

Abdominal muscle activation increases lumbar spinal stability: analysis of contributions of different muscle groups.

BACKGROUND: Antagonistic activation of abdominal muscles and increased intra-abdominal pressure are associated with both spinal unloading and spinal stabilization. Rehabilitation regimens have been proposed to improve spinal stability via selective recruitment of certain trunk muscle groups. This biomechanical analytical study addressed whether lumbar spinal stability is increased by such selective activation. METHODS: The biomechanical model included anatomically realistic three-layers of curved abdominal musculature, rectus abdominis and 77 symmetrical pairs of dorsal muscles. The muscle activations were calculated with the model loaded with either flexion, extension, lateral bending or axial rotation moments up to 60 Nm, along with intra-abdominal pressure up to 5 or 10 kPa (37.5 or 75 mm Hg) and partial bodyweight. After solving for muscle forces, a buckling analysis quantified spinal stability. Subsequently, different patterns of muscle activation were studied by forcing activation of selected abdominal muscles to at least 10% or 20% of maximum. FINDINGS: Spinal stability increased by an average factor of 1.8 with doubling of intra-abdominal pressure. Forcing at least 10% activation of obliques or transversus abdominis muscles increased stability slightly for efforts other than flexion, but forcing at least 20% activation generally did not produce further increase in stability. Forced activation of rectus abdominis did not increase stability. INTERPRETATION: Based on analytical predictions, the degree of stability was not substantially influenced by selective forcing of muscle activation. This casts doubt on the supposed mechanism of action of specific abdominal muscle exercise regimens that have been proposed for low back pain rehabilitation.

Refinement of elastic, poroelastic, and osmotic tissue properties of intervertebral disks to analyze behavior in compression.

Intervertebral disks support compressive forces because of their elastic stiffness as well as the fluid pressures resulting from poroelasticity and the osmotic (swelling) effects. Analytical methods can quantify the relative contributions, but only if correct material properties are used. To identify appropriate tissue properties, an experimental study and finite element analytical simulation of poroelastic and osmotic behavior of intervertebral disks were combined to refine published values of disk and endplate properties to optimize model fit to experimental data. Experimentally, nine human intervertebral disks with adjacent hemi-vertebrae were immersed sequentially in saline baths having concentrations of 0.015, 0.15, and 1.5 M and the loss of compressive force at constant height (force relaxation) was recorded over several hours after equilibration to a 300-N compressive force. Amplitude and time constant terms in exponential force-time curve-fits for experimental and finite element analytical simulations were compared. These experiments and finite element analyses provided data dependent on poroelastic and osmotic properties of the disk tissues. The sensitivities of the model to alterations in tissue material properties were used to obtain refined values of five key material parameters. The relaxation of the force in the three bath concentrations was exponential in form, expressed as mean compressive force loss of 48.7, 55.0, and 140 N, respectively, with time constants of 1.73, 2.78, and 3.40 h. This behavior was analytically well represented by a model having poroelastic and osmotic tissue properties with published tissue properties adjusted by multiplying factors between 0.55 and 2.6. Force relaxation and time constants from the analytical simulations were most sensitive to values of fixed charge density and endplate porosity.

Nonfusion treatment of adolescent idiopathic scoliosis by growth modulation and remodeling.

BACKGROUND: Adolescent idiopathic scoliosis (AIS) is a common disorder in which the spine gradually develops a curvature that is first detected in patients between 11 and 17 years of age. The only accepted treatment methods are bracing and surgery. Whether brace treatment alters the natural history is being questioned, and patient compliance is low. Surgery usually includes a spinal fusion that creates a rigid spine and concentrates stresses at the ends. METHODS: This study focuses on correlating the laboratory results with clinical reports for treating patients with AIS. In the laboratory, scoliosis with vertebral wedging has been created by asymmetric mechanical loading and has been corrected by reversing the loading. In the clinic, bracing and derotational casting have been successful in some reports, but compliance has been a problem with bracing and derotational casts have mainly been used in young children. Operative treatment has been successful, but a nonfusion operation remains elusive. FINDINGS AND RESULTS: In the laboratory, axial loading of growth plates altered growth according to the Hueter-Volkmann law, which states that compression decreases and distraction increases growth. Asymmetric loading of the spine caused asymmetric growth resulting in scoliosis with vertebral wedging. Asymmetric loading of tail vertebrae created vertebral wedging according to Wolff's law, which states that the bone remodels over time in response to prevailing mechanical demands. In the clinic, studies have shown that bracing may work if patients wore the brace as prescribed. Derotational casting in young children has been shown to prevent progression and even correct the scoliosis in some patients. Convex vertebral stapling has been successful in mild curves, but the results in larger curves have been disappointing. Anterolateral tethering has been successful in mild curves in young patients, but there is limited experience with this technique in patients with large curves. CONCLUSIONS: A brace that applies the appropriate loading and is worn as prescribed may dramatically improve the results of brace treatment. A procedure using external fixation or adjustable anterolateral tethering may achieve a nonfusion correction of AIS. LEVEL OF EVIDENCE: Level II.

Terminology - glossary including acronyms and quotations in use for the conservative spinal deformities treatment: 8th SOSORT consensus paper.

BACKGROUND: This report is the SOSORT Consensus Paper on Terminology for use in the treatment of conservative spinal deformities. Figures are provided and relevant literature is cited where appropriate. METHODS: The Delphi method was used to reach a preliminary consensus before the meeting, where the terms that still needed further clarification were discussed. RESULTS: A final agreement was found for all the terms, which now constitute the base of this glossary. New terms will be added after being discussed and accepted. DISCUSSION: When only one set of terms is used for communication in a place or among a group of people, then everyone can clearly and efficiently communicate. This principle applies for any professional group. Until now, no common set of terms was available in the field of the conservative treatment of scoliosis and spinal deformities. This glossary gives a common base language to draw from to discuss data, findings and treatment.

Intra-abdominal pressure and abdominal wall muscular function: Spinal unloading mechanism.

BACKGROUND: The roles of antagonistic activation of abdominal muscles and of intra-abdominal pressurization remain enigmatic, but are thought to be associated with both spinal unloading and spinal stabilization in activities such as lifting. Biomechanical analyses are needed to understand the function of intra-abdominal pressurization because of the anatomical and physiological complexity, but prior analyses have been over-simplified. METHODS: To test whether increased intra-abdominal pressure was associated with reduced spinal compression forces for efforts that generated moments about each of the principal axis directions, a previously published biomechanical model of the spine and its musculature was modified by the addition of anatomically realistic three-layers of curved abdominal musculature connected by fascia to the spine. Published values of muscle cross-sectional areas and the active and passive stiffness properties were assigned. The muscle activations were calculated assuming minimized muscle stress and stretch for the model loaded with flexion, extension, lateral bending and axial rotation moments of up to 60 Nm, along with intra-abdominal pressurization of 5 or 10 kPa (37.5 or 75 mm Hg) and partial bodyweight (340 N). FINDINGS: The analysis predicted a reduction in spinal compressive force with increase in intra-abdominal pressurization from 5 to 10 kPa. This reduction at 60 Nm external effort was 21% for extension effort, 18% for flexion effort, 29% for lateral bending and 31% for axial rotation. INTERPRETATION: This analysis predicts that intra-abdominal pressure produces spinal unloading, and shows likely muscle activation patterns that achieve this.

The role of remodeling and asymmetric growth in vertebral wedging.

BACKGROUND: Scoliosis with vertebral wedging is thought to be caused by asymmetric growth (Hueter-Volkmann law), but vertebral diaphyseal remodeling (Wolff's law) may also contribute to the deformity. We investigated whether vertebral wedging in scoliosis might involve both mechanisms. METHODS: An external fixator was used to impose a 30 degrees scoliosis and compression of 0.1 or 0.2 MPa to the tails of 10 5-week-old and 20 14-week-old Sprague-Dawley rats for 6 weeks. The rats were divided into three groups of 10 animals each: Group 1: 5-week-old animals with 0.1 MPa compression; Group 2: 14-week-old animals with 0.1 MPa compression; Group 3: 14-week-old animals with 0.2 MPa compression. Vertebral wedging and diaphyseal curvature were measured from micro CT scans performed at weeks 1, 3, and 6. Wedging due to asymmetrical growth and remodeling was calculated from a Calcein label administered at week 3 and a Xylenol label at week 6. RESULTS: The growth rate of the loaded vertebrae as a per cent of control vertebrae was 60% in Group 1, 40% in Group 2, and 30% in Group 3. The growth rate of control vertebrae in 14-week-old animals was 16% that of 5-week-old animals. The animals in all 3 groups developed a scoliosis with vertebral wedging that averaged 18.7 degrees in Group 1, 8.2 degrees in Group 2, and 10.1 degrees in Group 3. Asymmetric growth was much greater in Group 1 (5-week-old) animals. The ossified epiphyses became wedged and diaphyseal remodeling occurred in all groups. CONCLUSIONS: The major contribution to the vertebral wedging was asymmetric growth in the 5-week-old animals and diaphyseal remodeling in the 14-week-old animals. The results support the concept that if appropriate loads can be applied to human vertebrae through minimally invasive techniques, scoliosis and vertebral wedging can be corrected without a spinal fusion in both adolescents and adults.