Load-shift--numerical evaluation of a new design philosophy for uncemented hip prostheses

All hip replacement prostheses alter the load transfer from the hip joint into the femur by changing the mechanical loading of the proximal femur from an externally to an internally loaded system. This alteration of the load transfer causes stress shielding and might lead to severe bone loss, especially with uncemented prostheses. To minimize these effects, load transfer to the femur should occur as proximal as possible. In order to support sufficient primary stability, however, directly post operative (PO) distal stabilization is reasonable. Consequently, the prostheses have to alter its mechanical characteristics after implantation. This concept is referred to as load-shift concept. Primary stability during the early PO state is provided by a prosthesis shaft, which is widened at the tip by a biodegradable pin. After resorption of the pin load transfer occurs no longer distally. The objective of this study was the numerical evaluation of the load-shift concept. The analysis was performed with a finite element model. Three-dimensional non-linear dynamic gait analyses data were used to evaluate whether the load transfer during walking can be altered effectively by insertion and resorption of a distal pin. Directly PO 38% of the transverse load is transferred through the prosthesis shaft and micromotion of the proximal prostheses tip is below 55 microm. After resorption of the pin, no transverse loads are transferred through the prosthesis shaft. Therefore, the loading of the proximal bone tissue is far more pronounced than in the case of a standard prosthesis, demonstrating the feasibility of the load-shift concept. A balanced degradation of the pin simultaneously with the ingrowth of the prosthesis is expected to reduce hip replacement complications.     

3 种不同弹性模量钛合金股骨假体在羊股骨置换模型中应力分布的三维 有限元分析

目的:通过三维有限元分析方法来观察并比较3种不同弹性模量钛合金股骨假体在羊股骨置换模型中von-Mises应力分布的情况。方法:采用64排螺旋CT对一健康成年羊的下肢股骨进行全长的CT扫描,扫描层厚为0.5 mm,扫描所得的数据存储为DICOM文件。将得到的DICOM文件导入到CT图像分析软件Mimics 10.0,然后利用Mimics 10.0软件来生成股骨的骨质点云数据,再将生成的骨质点云数据导入到Simpleware分析软件,通过机械加工反求中的复杂曲面造型技术建立起精确的三维实体模型。对三维实体模型进行网格划分,确定了髓腔的形状,并根据羊下肢股骨髓腔的形状设计了作者实验用的羊股骨假体模型,然后在ANSYS 12.1软件中进行网格划分。给予加载缓慢行走载荷以及扭转载荷,分析并比较羊股骨以及3种不同弹性模量钛合金股骨假体在股骨置换模型中von-Mises应力分布的情况。结果:在缓慢行走载荷以及扭转载荷条件下,3种不同弹性模量钛合金股骨假体von-Mises应力分布变化趋势一致,假体的柄颈结合部以及假体柄上1/3为应力集中区域。3种不同弹性模量的最大应力集中点均位于柄颈结合部,60 GPa弹性模量的股骨假体植入后假体的最大应力最小(37.8 MPa、29.1 MPa),股骨的最大应力最大(12.6 MPa、24.5 MPa);80 GPa的次之,假体的最大应力(38.4 MPa、33.4 MPa),股骨的最大应力(12.5 MPa、24.5 MPa);110 GPa的股骨假体植入后假体的最大应力最大(38.9 MPa、38.1 MPa),股骨的最大应力最小(12.3 MPa、24.5 MPa)。60 GPa弹性模量的股骨假体植入后的假体最大位移和相对位移均最小(缓慢行走载荷下假体最大位移为0.551 mm、相对位移为0.008 mm,扭转载荷下假体最大位移为0.730 mm、相对位移为0.011 mm)。结论:较低弹性模量的钛合金股骨假体(60 GPa)由于其弹性模量更接近于骨组织的弹性模量,股骨假体与股骨间的"应力遮挡"效应较小,更有利于应力在股骨假体及股骨间的传递,增加了股骨假体的早期稳定性,延长了其临床寿命。     

Artificial composite bone as a model of human trabecular bone: the implant-bone interface

The use of artificial bones in implant testing has become popular due to their low variability and ready availability. However, friction coefficients, which are critical to load transfer in uncemented implants, have rarely been compared between human and artificial bone, particularly for wet and dry conditions. In this study, the static and dynamic friction coefficients for four commercially used titanium surfaces (polished, Al(2)O(3) blasted, plasma sprayed, beaded) acting on the trabecular component of artificial bones (Sawbones) were compared to those for human trabecular bone. Artificial bones were tested in dry and wet conditions and normal interface stress was varied (0.25, 0.5, 1.0MPa). Friction coefficients were mostly lower for artificial bones than real bone. In particular, static friction coefficients for the dry polished surface were 20% of those for real bone and 42-61% for the dry beaded surface, with statistical significance (alpha<0.05). Less marked differences were observed for dynamic friction coefficients. Significant but non-systematic effects of normal stress or wet/dry condition on friction coefficients were observed within each surface type. These results indicate that the use of artificial bone models for pre-clinical implant testing that rely on interface load transfer with trabecular bone for mechanical integrity can be particularly sensitive to surface finish and lubrication conditions.     

The dynamic flexion/extension properties of the lumbar spine in vitro using a novel pendulum system

The biomechanical properties of the ligamentous cadaver spine have been previously examined using a variety of experimental testing protocols. Ongoing technical challenges in the biomechanical testing of the spine include the application of physiologic compressive loads and the application of dynamic bending moments while allowing unconstrained three-dimensional motion. The purpose of this study was to report the development of a novel pendulum apparatus that addressed these challenges and to determine the effects of various axial compressive loads on the dynamic biomechanical properties of the lumbar functional spinal unit (FSU). Lumbar FSUs were tested in flexion and extension under five axial compressive loads chosen to represent physiologic loading conditions. After an initial rotation, the FSUs behaved as a dynamic, underdamped vibrating elastic system. Bending stiffness and coefficient of damping increased significantly as the compressive pendulum load increased. The apparatus described herein is a relatively simple approach to determining the dynamic bending properties of the FSU, and potentially disc arthroplasty devices. It is capable of applying physiologic compressive loads at dynamic rates without constraining the kinematics of the joints, crucial requirements for testing FSUs in vitro.     

A trial of evaluating static and dynamic loads in man by means of analysis of the variability of instantaneous heart rate

The instantaneous heart rate changes were examined in 12 forest workers during following tests: increasing (up to refusal) dynamic load on ergometer, rest in erect position, static load (about 75% of body weight) in erect position. The ECG signal, carrying information on heart rate, was amplified and processed by means of an especially designed heart rate analyzer. The dynamics of heart rate changes was evaluated through the analysis of differences of subsequent instantaneous heart rates and of distributions of heart rate accelerations and decelerations during the tests. Significantly different distributions of instantaneous heart rate changes as well as of accelerations and decelerations were observed for different loads.     

Quantification of structural loading during off-road cycling.

To provide data for fatigue life prediction and testing of structural components in off-road bicycles, the objective of the research described herein was to quantify the loads input to an off-road bicycle as a result of surface-induced loads. A fully instrumented test bicycle was equipped with dynamometers at the pedals, handlebars, and hubs to measure all in-plane structural loads acting through points of contact between the bicycle and both the rider and the ground. A portable data acquisition system carried by the standing rider allowed, for the first time, this loading information to be collected during extended off-road testing. In all, seven experienced riders rode a downhill trial test section with the test bicycle in both front-suspension and full-suspension configurations. The load histories were used quantitatively to describe the load components through the computation of means, standard deviations, amplitude probability density functions, and power spectral density functions. For the standing position, the coefficients of variation for the load components normal to the ground were greater than 1.2 for handlebar forces and 0.3 and 0.5-0.6 for the pedal and hub forces, respectively. Thus, the relative contribution of the dynamic loading was much greater than the static loading at the handlebars but less so at the pedals and hubs. As indicated by the rainflow count, high amplitude loading was developed approaching 3 and 5 times the weight of the test subjects at the front and rear wheels, respectively. The power spectral densities showed that energy was concentrated in the band 0-50 Hz. Through stress computations and knowledge of material properties, the data can be used analytically to predict the fatigue life of important structural components such as those for steering. The data can also be used to develop a fatigue testing protocol for verifying analytical predictions of fatigue life.     

Strength requirements for internal and external prostheses

Throughout the history of development of joint replacement implants and external prostheses there have been mechanical failures due to a discrepancy between material strength, cross-sectional characteristics and the loads developed in normal or abnormal function by the patient utilising the device. Particularly for internal prostheses attention is being paid at the present time to wear characteristics and the requirements for the articulating surfaces and the volume of wear particles produced during tests simulating the use of the device within the patient. The particular importance of the wear particles is that they seem to be associated with accelerated resorption of bone at areas essential for successful fixation of the implant within it. This article will consider joint replacements at the knee and hip and external prostheses for the leg. If failure due to external trauma is ignored the loads to be considered in testing standards correspond in implants to the muscular and ligamentous forces related to the forces developed between ground and foot and to the bending moments in the structure of leg prostheses. Generally it can be assumed that the treatment of the patient following trauma is more easily accomplished and more likely to be successful if the prosthesis has failed and not the bony structure of the patient. However, the author is unaware that these devices have ever been designed to have lower intrinsic strength than the anatomical structures to which they are connected; indeed in many cases particularly for implants they are much stronger than the bone to which they are connected. The major difficulty in rational design of prosthetic devices has been uncertainty about the importance of occasionally applied loads of a high value relative to those on a frequent basis and also to the frequency of application of these overloads. In this paper consideration is given to methods of determination of load systems relevant to the mechanical performance of implanted joint replacements at the hip and the knee and external prostheses for leg amputees. New data are presented relating to walking, other daily activities and the corresponding frequency of occurrence of these. Loading data on implants obtained by various biomechanical models is compared and related to the loads actually measured by implanted transducers. The philosophy of the standardised test load systems and the performance requirements is reviewed.     

An experimental method for measuring force on the spinal facet joint: description and application of the method.

A technique is described for measuring load magnitude and resultant load contact location in the facet joint in response to applied loads and moments, and the technique applied to the canine lumbar spine motion segment. Due to the cantilever beam geometry of the cranial articular process, facet joint loads result in surface strains on the lateral aspect of the cranial articular process. Strains were quantified by four strain gages cemented to the bony surface of the process. Strain measured at any one gage depended on the loading site on the articular surface of the caudal facet and on the magnitude of the facet load. Determination of facet loads during in vitro motion segment testing required calibration of the strains to known loads of various magnitudes applied to multiple sites on the caudal facet. The technique is described in detail, including placement of the strain gages. There is good repeatability of strains to applied facet loads and the strains appear independent of load distribution area. Error in the technique depends on the location of the applied facet loads, but is only significant in nonphysiologic locations. The technique was validated by two independent methods in axial torsion. Application of the technique to five in vitro canine L2-3 motion segments testing resulted in facet loads (in newtons, N) of 74+ / -23 N (mean + / -STD) in 2 newton-meter, Nm, extension, to unloaded in flexion. Lateral bending resulted in loads in the right facet of 40+ / -32 N for 1 Nm right lateral bending and 54+ / -29 N for 1 Nm left lateral bending. 4 Nm Torsion with and without 100 N axial compression resulted in facet loads of 92+ / -27 N and 69+ / -19 N, respectively. The technique is applicable to dynamic and in vivo studies.     

A new technique to measure the dynamic contact pressures on the Tibial Plateau

Studies of the load transfer role of the meniscus have been limited to static experimental and analytical approaches. The objective of this study was to develop an experimental technique to allow the contact pressures on the tibial plateau of cadaveric knees to be measured under dynamic physiological loads. Accordingly, we adapted a load-controlled knee joint simulator to accept a cadaveric sheep knee, programmed the simulator with sheep gait kinematics data, and utilized a pressure sensor array to measure the contact pressure distribution on the lateral tibial plateau during gait. The technique was applied to six sheep knees that were tested intact and after meniscectomy. Meniscectomy resulted in a 267% increase in average contact pressure, a 117% increase in peak contact pressure, and an 80% decrease in contact area, all measured at the point of maximum peak contact stress in the gait cycle. It is envisaged that the experimental model herein developed will allow for the screening of candidate materials prior to more expensive and time-consuming animal models.     

Effects of loading on the biochemical behavior of molars of Homo,Pan, and Pongo

In a previous study, we found systematic differences in the biomechanical behavior of modern human molars using finite element stress analyses (FESA), which led us to propose that molars are adapted to differently-directed loads depending on their position within the mouth (Spears and Macho [1998] Am. J. Phys. Anthropol. 106:467–482 ). While the FESA results thus derived have not been verified experimentally, such an interpretation seemed reasonable. To refine the model previously presented, this study assessed the effects of 1) food particle size on the biomechanical behavior of molars, and those of 2) differences in morphology, particularly enamel thickness, on stress distribution. In order to appraise the evolutionary significance of the findings, the FESA results for modern humans were subsequently compared with those obtained for molars of one individual of Pan and Pongo, respectively. Bearing in mind limitations imposed by the FESA models created and analyzed in this study, constant cleavage-type loads and cuspal tip loads at different directions were employed on all teeth: this facilitated comparisons of patterns of stress distribution across molars and species. In Pan and Homo, cleavage-type loads exerted by big food particles tended to be better dissipated anteriorly than posteriorly, although trends in Pongo were less clear-cut. Furthermore, similar to modern humans, the buccal cusps of mandibular molars appeared to be able to dissipate the loads associated with a pestle-type action, while maxillary molars were better designed to dissipate the loads which would result if they acted as mortars against which the food is crushed/ground. While increases in enamel thickness lowered the overall stress values in teeth only slightly, changes in outer morphology could have a more profound effect on these stress levels. Overall, Pan appeared to be most generalized, while Homo and Pongo showed a number of unique specializations, which are in accordance with what is currently understood about their respective masticatory apparatus and dietary niche. Am J Phys Anthropol 109:211–227, 1999.© 1999 Wiley-Liss, Inc.     

Effect of load position on muscle forces, internal loads and stability of the human spine in upright postures

A novel kinematics-based approach coupled with a non-linear finite element model was used to investigate the effect of changes in the load position and posture on muscle activity, internal loads and stability margin of the human spine in upright standing postures. In addition to 397 N gravity, external loads of 195 and 380 N were considered at different lever arms and heights. Muscle forces, internal loads and stability margin substantially increased as loads displaced anteriorly away from the body. Under same load magnitude and location, adopting a kyphotic posture as compared with a lordotic one increased muscle forces, internal loads and stability margin. An increase in the height of a load held at a fixed lever arm substantially diminished system stability thus requiring additional muscle activations to maintain the same margin of stability. Results suggest the importance of the load position and lumbar posture in spinal biomechanics during various manual material handling operations.     

Reduced nucleus pulposus glycosaminoglycan content alters intervertebral disc dynamic viscoelastic mechanics

The intervertebral disc functions over a range of dynamic loading regimes including axial loads applied across a spectrum of frequencies at varying compressive loads. Biochemical changes occurring in early degeneration, including reduced nucleus pulposus glycosaminoglycan content, may alter disc mechanical behavior and thus may contribute to the progression of degeneration. The objective of this study was to determine disc dynamic viscoelastic properties under several equilibrium loads and loading frequencies, and further, to determine how reduced nucleus glycosaminoglycan content alters dynamic mechanics. We hypothesized that (1) dynamic stiffness would be elevated with increasing equilibrium load and increasing frequency, (2) the disc would behave more elastically at higher frequencies, and finally, (3) dynamic stiffness would be reduced at low equilibrium loads under all frequencies due to nucleus glycosaminoglycan loss. We mechanically tested control and chondroitinase ABC injected rat lumbar motion segments at several equilibrium loads using oscillatory loading at frequencies ranging from 0.05 to 5 Hz. The rat lumbar disc behaved non-linearly with higher dynamic stiffness at elevated compressive loads irrespective of frequency. Phase angle was not affected by equilibrium load, although it decreased as frequency was increased. Reduced glycosaminoglycan decreased dynamic stiffness at low loads but not at high equilibrium loads and led to increased phase angle at all loads and frequencies. The findings of this study demonstrate the effect of equilibrium load and loading frequencies on dynamic disc mechanics and indicate possible mechanical mechanisms through which disc degeneration can progress.     

Modeling the biomechanics of the mandible: a three-dimensional finite element study.

Three-dimensional finite element models of a partially edentulated human mandible were generated to calculate the mechanical response to simulated isometric biting and mastication loads. The level of mesh refinement was established via a convergence test and showed that a model with over 30,000 degrees of freedom was required to obtain analysis accuracy. The functional loading cases included muscle loading based on an algorithm that assigns muscle forces in accordance with muscle cross-sectional area, while maintaining static equilibrium. Results were found for isometric application of unilateral and bilateral bite and mastication loading, and two different sets of displacement boundary conditions were imposed at the condyles. The mechanical response is shown in terms of displacements, principal strains, and a new measure called the 'mechanical intensity scalar'. For each load case studied, there was substantial bending in the molar region of the corpus and high tensile strains in the anterior portion of the ramus.     

Loosening of cementless femoral stems: a biomechanical analysis of immediate fixation with loading vertical, femur horizontal

Large joint implants must have immediate fixation to be successful. Unfortunately, the magnitude and consistency of achieving this remains largely unknown. For cementless femoral components it is being increasingly appreciated that torsional loading as occurs during stair climbing or rising from a chair leads to loosening and thigh pain in some cases. A biomechanical test was developed to evaluate fixation in this position. Twelve pairs of human cadaveric femora were press-fit with an AML stem. Each femur was secured in a horizontal position, and the prosthetic head cyclically loaded in a vertically downward direction. The offset of the prosthetic head resulted in a combined torsional and compressive load being applied to the stem within the proximal femur. Loosening was found to consistently occur and rapidly accelerate when the head subsided more than 0.2 mm during 100 cycles. For the AML stem, loosening developed at loads from 62 to 171% of body weight and after as few as 800 cycles. This is within the physiologic range of normal daily activities as measured by others with instrumented prostheses. This poses a challenge to the ability of press-fit stems to tolerate torsional loads in vivo. Patients with a cementless prosthesis should be protected from torsional loading until porous ingrowth and/or bone remodelling have had time to occur. Testing the same stem in paired femora demonstrated no right vs left difference (p greater than 0.6).     

Effect of load position on muscle forces,internal loads and stability of the human spine in upright postures

A novel kinematics-based approach coupled with a non-linear finite element model was used to investigate the effect of changes in the load position and posture on muscle activity, internal loads and stability margin of the human spine in upright standing postures. In addition to 397 N gravity, external loads of 195 and 380 N were considered at different lever arms and heights. Muscle forces, internal loads and stability margin substantially increased as loads displaced anteriorly away from the body. Under same load magnitude and location, adopting a kyphotic posture as compared with a lordotic one increased muscle forces, internal loads and stability margin. An increase in the height of a load held at a fixed lever arm substantially diminished system stability thus requiring additional muscle activations to maintain the same margin of stability. Results suggest the importance of the load position and lumbar posture in spinal biomechanics during various manual material handling operations.     

Regulating explosive resistance training intensity using the rating of perceived exertion

Explosive resistance training (ERT) improves older adults' strength and power, and methods to make this form of training more accessible and useful to older adults are needed. The purpose of this study was to evaluate whether the rating of perceived exertion (RPE) scale would predict a broad range of ERT intensities on the leg press with older adults. If successful, then a load-RPE relationship could be used to regulate the intensity of training loads for ERT with older adults, allowing the elimination of maximal strength testing. Twenty-one older adults (≥65 years) with resistance training experience took part in 2 testing sessions. Session 1: Subjects performed high-velocity repetitions on the leg press for up to 9 loads (from 60 to 140% body weight) presented in quasi-randomized order, and then reported their RPE for each load. Session 2: A 1 repetition maximum (1RM) strength test was conducted. Regression analysis revealed that the average RPE across subjects for each load strongly predicted the average %1RM across subjects (R2 = 99.5%; p < 0.001). This allows the establishment of a load-RPE relationship for use in selecting ERT loads for older adults on the leg press. For example, high-intensity loads (70-90% 1RM) that would elicit both strength and power gains when used with ERT aligned with an RPE of 14-16. Lighter loads that may be useful for training for power, but not strength (<70% 1RM), were identified with RPE scores of 13 and lower. The load-RPE relationship may simplify the regulation of intensity of ERT with older adults on the leg press, where the exercising older adult could be guided to select loads according to their RPE.     

Peak force and rate of force development during isometric and dynamic mid-thigh clean pulls performed at various intensities

Eight male collegiate weightlifters (age: 21.2 +/- 0.9 years; height: 177.6 +/- 2.3 cm; and body mass: 85.1 +/- 3.3 kg) participated in this study to compare isometric to dynamic force-time dependent variables. Subjects performed the isometric and dynamic mid-thigh clean pulls at 30-120% of their one repetition maximum (1RM) power clean (118.4 +/- 5.5 kg) on a 61 x 121.9-cm AMTI forceplate. Variables such as peak force (PF) and peak rate of force development (PRFD) were calculated and were compared between isometric and dynamic conditions. The relationships between force-time dependent variables and vertical jump performances also were examined. The data indicate that the isometric PF had no significant correlations with the dynamic PF against light loads. On the one hand, there was a general trend toward stronger relationships between the isometric and dynamic PF as the external load increased for dynamic muscle actions. On the other hand, the isometric and dynamic PRFD had no significant correlations regardless of the external load used for dynamic testing. In addition, the isometric PF and dynamic PRFD were shown to be strongly correlated with vertical jump performances, whereas the isometric PRFD and dynamic PF had no significant correlations with vertical jump performances. In conclusion, it appears that the isometric and dynamic measures of force-time curve characteristics represent relatively specific qualities, especially when dynamic testing involves small external loads. Additionally, the results suggest that athletes who possess greater isometric maximum strength and dynamic explosive strength tend to be able to jump higher.     

Evaluation of cement stresses in finite element analyses of cemented orthopaedic implants.

Stress analysis of the cement fixation of orthopaedic implants to bone is frequently carried out using finite element analysis. However the stress distribution in the cement layer is usually intricate, and it is difficult to report it in a way that facilitates comparison of implants for pre-clinical testing. To study this problem, and make recommendations for stress reporting, a finite element analysis of a hip prosthesis implanted into a synthetic composite femur is developed. Three cases are analyzed: a fully bonded implant, a debonded implant, and a debonded implant where the cement is removed distal to the stem tip. In addition to peak stresses, and contour and vector plots, a stressed volume and probability-of-failure analysis is reported. It is predicted that the peak stress is highest for the debonded stem, and that removal of the distal cement more than halves this peak stress. This would suggest that omission of the distal cement is good for polished prostheses (as practiced for the Exeter design). However, if the percentage of cement stressed above a certain threshold (say 3 MPa) is considered, then the removal of distal cement is shown to be disadvantageous because a higher volume of cement is stressed to above the threshold. Vector plots clearly demonstrate the different load transfer for bonded and debonded prostheses: A bonded stem generates maximum tensile stresses in the longitudinal direction, whereas a debonded stem generates most tensile stresses in the hoop direction, except near the tip where tensile longitudinal stresses occur due to subsidence of the stem. Removal of the cement distal to the tip allows greater subsidence but alleviates these large stresses at the tip, albeit at the expense of increased hoop stresses throughout the mantle. It is concluded that a thorough analysis of cemented implants should not report peak stress, which can be misleading, but rather stressed volume, and that vector plots should be reported if a precise analysis of the load transfer mechanism is required.     

A Computational Model of a Descending Mechanosensory Pathway Involved in Active Tactile Sensing

Many animals, including humans, rely on active tactile sensing to explore the environment and negotiate obstacles, especially in the dark. Here, we model a descending neural pathway that mediates short-latency proprioceptive information from a tactile sensor on the head to thoracic neural networks. We studied the nocturnal stick insect Carausius morosus, a model organism for the study of adaptive locomotion, including tactually mediated reaching movements. Like mammals, insects need to move their tactile sensors for probing the environment. Cues about sensor position and motion are therefore crucial for the spatial localization of tactile contacts and the coordination of fast, adaptive motor responses. Our model explains how proprioceptive information about motion and position of the antennae, the main tactile sensors in insects, can be encoded by a single type of mechanosensory afferents. Moreover, it explains how this information is integrated and mediated to thoracic neural networks by a diverse population of descending interneurons (DINs). First, we quantified responses of a DIN population to changes in antennal position, motion and direction of movement. Using principal component (PC) analysis, we find that only two PCs account for a large fraction of the variance in the DIN response properties. We call the two-dimensional space spanned by these PCs ‘coding-space’ because it captures essential features of the entire DIN population. Second, we model the mechanoreceptive input elements of this descending pathway, a population of proprioceptive mechanosensory hairs monitoring deflection of the antennal joints. Finally, we propose a computational framework that can model the response properties of all important DIN types, using the hair field model as its only input. This DIN model is validated by comparison of tuning characteristics, and by mapping the modelled neurons into the two-dimensional coding-space of the real DIN population. This reveals the versatility of the framework for modelling a complete descending neural pathway.     

POLLINATION ADAPTATION IN PEDICULARIS GROENLANDICA

Cinematographic and stereophotographic records indicate that Pedicularis groenlandica is pollinated in the Colorado Front Range by seven species of pollen-foraging bumblebees (Bombus sp.) to which the nectarless flower is intricately adapted functionally and structurally. Removing pollen by wing vibrations of an oscilloscopically identified frequency significantly distinct from flight vibration frequency, foragers carried pollen loads with up to three foreign pollen types in addition to Pedicularis pollen, which was found in all loads. No direct correlation was evident between flight vibration frequency and combined body-pollen load weight, ambient air temperature, or forager species. The flower is phenologically and morphologically adapted to the worker caste of apparently any Bombus species available to it throughout the plant's montane-alpine zone vertical-distribution range. The evolutionary position of the pollination mechanism is considered in relation to the floral morphology of other species in the genus Pedicularis.