Biomechanical adaptation of ulnar cross-sectional morphology in brachiating primates

The forelimbs of hylobatids (gibbons and siamang) are distinctive among tetrapods in that they are loaded in overall tension during normal locomotion. While hylobatid ulnae must also encounter bending stresses in the course of their full range of locomotor behavior, their loading regime differs from that of quadrupedal anthropoids in that these bending stresses are distributed evenly along the bone, are not exerted in a preferred plane, and are probably of generally lower magnitude. This study examines the degree to which hylobatid ulnae are adapted to this suspensory loading regime. We obtained cross-sections of ulnae at various increments along the length of the bone using CAT scans. The sample comprises 476 cross-sections representing the ulnae of 25 individuals from five species of comparable body size. We show that in gibbons and siamang, the patterning of ulnar cross-sectional area and resistance to bending in the dorsoventral plane along the ulnar diaphysis differ from that of similarly sized quadrupedal anthropoids in the manner predicted by a suspensory loading regime. We also find the same pattern for the ulnae of Ateles, whose loading regime may be fairly similar to that of hylobatids. However, we find that the cross-sectional shape of the ulnar diaphysis in hylobatids and Ateles does not differ from that of quadrupedal monkeys in the manner predicted by a suspensory loading regime. © 1995 Wiley-Liss, Inc.     

Site specific bone adaptation response to mechanical loading

Over 25 million Americans suffer from osteoporosis. Bone size and strength depends both upon the level of adaptation due to physical activity (applied load), and genetics. We hypothesized that bone adaptation to loads differs among mice breeds and bone sites. Forty-five adult female mice from three inbred strains (C57BL/6 [B6], C3H/HeJ [C3], and DBA/2J [D2]) were loaded at the right tibia and ulna in vivo with non-invasive loading devices. Each loading session consisted of 99 cycles at a force range that induced approximately 2000 microstrain (microepsilon) at the mid-shaft of the tibia (2.5 to 3.5 N force) and ulna (1.5 to 2 N force). The right and left ulnae and tibiae were collected and processed using protocols for histological undecalcified cortical bone slides. Standard histomorphometry techniques were used to quantify new bone formation. The histomorphometric variables include percentage mineralizing surface (%MS), mineral apposition rate (MAR), and bone formation rate (BFR). Net loading response [right-left limb] was compared between different breeds at tibial and ulnar sites using two-way ANOVA with repeated measures (p<0.05). Significant site differences in bone adaptation response were present within each breed (p<0.005). In all the three breeds, the tibiae showed greater percentage MS, MAR and BFR than the ulna at similar in vivo load or mechanical stimulus (strain). These data suggest that the bone formation due to loading is greater in the tibiae than the ulnae. Although, no significant breed-related differences were found in response to loading, the data show greater trends in tibial bone response in B6 mice as compared to D2 and C3 mice. Our data indicate that there are site-specific skeletal differences in bone adaptation response to similar mechanical stimulus.     

Patterns of strain in the macaque ulna during functional activity

In vivo bone strain experiments were performed on the ulnae of three female rhesus macaques to test how the bone deforms during locomotion. The null hypothesis was that, in an animal moving its limbs predominantly in sagittal planes, the ulna experiences anteroposterior bending. Three rosette strain gauges were attached around the circumference of the bone slightly distal to midshaft. They permit a complete characterization of the ulna's loading environment. Strains were recorded during walking and galloping activities. Principal strains and strain directions relative to the long axis of the bone were calculated for each gauge site. In all three animals, the lateral cortex experienced higher tensile than compressive principal strains during the stance phase of walking. Compressive strains predominated at the medial cortex of two animals (the gauge on this cortex of the third animal did not function). The posterior cortex was subject to lower strains; the nature of the strain was highly dependent on precise gauge position. The greater principal strains were aligned closely with the long axis of the bone in two animals, whereas they deviated up to 45° from the long axis in the third animal. A gait change from walk to gallop was recorded for one animal. It was not accompanied by an incremental change in strain magnitudes. Strains are at the low end of the range of strain magnitudes recorded for walking gaits of nonprimate mammals. The measured distribution of strains in the rhesus monkey ulna indicates that mediolateral bending, rather than anteroposterior bending, is the predominant loading regime, with the neutral axis of bending running from anterior and slightly medial to posterior and slightly lateral. A variable degree of torsion was superimposed over this bending regime. Ulnar mediolateral bending is apparently caused by a ground reaction force vector that passes medial to the forearm. The macaque ulna is not reinforced in the plane of bending. The lack of buttressing in the loaded plane and the somewhat counterintuitive bending direction recommend caution with regard to conventional interpretations of long bone cross-sectional geometry. Am J Phys Anthropol 106:87–100, 1998. © 1998 Wiley-Liss, Inc.     

Cellular accommodation and the response of bone to mechanical loading

Several mathematical rules by which bone adapts to mechanical loading have been proposed. Previous work focused mainly on negative feedback models, e.g., bone adapts to increased loading after a minimum strain effective (MES) threshold has been reached. The MES algorithm has numerous caveats, so we propose a different model, according to which bone adapts to changes in its mechanical environment based on the principle of cellular accommodation. With the new algorithm we presume that strain history is integrated into cellular memory so that the reference state for adaptation is constantly changing. To test this algorithm, an experiment was performed in which the ulnae of Sprague-Dawley rats were loaded in axial compression. The animals received loading for 15 weeks with progressively decreasing loads, increasing loads, or a constant load. The results showed the largest increases in geometry in the decreasing load group, followed by the constant load group. Bone formation rates (BFRs) were significantly greater in the decreasing load group during the first 2 weeks of the study as compared to all other groups (P<0.05). After the first few weeks of mechanical loading, the BFR in the loaded ulnae returned to the values of the nonloaded ulnae. These experimental results closely fit the predicted results of the cellular accommodation algorithm. After the initial weeks of loading, bone stopped responding so the degree of adaptation was proportional to the initial peak load magnitude.     

Drought response and changing mean sensitivity of European beech close to the dry distribution limit

European beech (Fagus sylvatica L.) covers a large area mainly in the colline and montane ranges in Europe, and a drier and warmer climate, as expected for the coming decades, is likely to alter its distribution. So far, an altitudinal shift has been projected using a variety of modelling approaches. However, we lack knowledge about the climatic and edaphic factors that control the growth and competitive behaviour of beech at its dry distribution limit. We applied and further developed dendroecological methods to study the drought response and sensitivity pattern of beech at sites with different moisture regimes. We compared three pairs of sites from different geographical regions near the dry distribution limit of beech in Switzerland, consisting of a dry and mesic site each. Radial growth differed between mesic and dry sites, in that average ring-width at mesic sites was around double the width at dry sites. For the whole study period (1930–2006), the sites with the lowest available soil water capacity (AWC) were found to respond most sensitively to drought. However, in recent years, sites with higher AWC have shown increasing drought sensitivity, i.e. they have responded even more strongly to drought than the dry sites. This change in sensitivity corresponds to a seasonal shift in drought response at mesic sites, with a change in the months showing significant drought response in all three studied regions compared with the past. Even though dry sites generally displayed a larger number of negative pointer years than mesic sites, it appears that the frequency of pointer years has increased at mesic sites, i.e. they have become more sensitive particularly in the last quarter of the twentieth century. Yet, the frequency of pointer years at the dry sites has remained fairly constant. These results indicate that beech trees near their dry distribution limit are adapted to extreme conditions already, while changes in the growth patterns of beech under mesic conditions have to be expected.     

A torsional strength comparison of vascularized and nonvascularized bone grafts

Comparisons of torsional strength are made on the ulnae from the forelegs of short haired hounds where a nonvascularized graft was performed on one leg and a vascularized graft performed on the other. By using the forelegs of a dog as the experimental model and microsurgical techniques, a vascularized bone segment was used to graft a five centimeter nonunion in one leg and at the same time a conventional bone graft was performed on a similar nonunion in the other leg. Similar segments of rib bone were used for each graft. Torsional strength data are shown for nine experimental animals. A successful method for mounting the bones for testing of torsional strength in a torsion machine is given. In each case for which the bones healed properly, the vascularized bone graft proved to be significantly stronger in torsion.     

Seasonal changes in biom ass of macrophytes on shaded and unshaded sections of the River Lambourn, England

SUMMARY. Monthly changes in the biomass of Ranunculus, Berula and Callitriche were recorded on two 50-m sites on the River Lambourn at Bagnor between March 1971 and October 1973. On the unshaded site. Ranunculus was the dominant macrophyte and characteristically its biomass changed from 20–40 g dry wt m^-2 in March to around 400 g dry wt m^-2 at the end of the growing season. On the shaded site, where Berula was the dominant macrophyte. the biomass of Ranunculus was lower than on the unshaded site. There was no significant difference between the mean biomasses of Berula on these two sites but Callitriche had a signifieantly higher biomass on the shaded site. Estimates of the total biomass of macrophyte on each site were calculated and the biomass of macrophyte removed during cutting operations each summer was also assessed.     

Variation of trabecular architecture in proximal femur of postmenopausal women

This investigation of microstructure in the human proximal femur probes the relationship between the parameters of the FRAX index of fracture risk and the parameters of bone microstructure. The specificity of fracture sites at the proximal femur raises the question of whether trabecular parameters are site-specific during post-menopause, before occurrence of fragility fracture. The donated proximal femurs of sixteen post-menopausal women in the sixth and seventh decades of life, free of metabolic pathologies and therapeutic interventions that could have altered the bone tissue, constituted the material of the study. We assessed bone mineral density of the proximal femurs by dual energy X-ray absorptiometry and then sectioned the femurs through the center of the femoral head and along the femoral neck axis. For each proximal femur, morphometry of trabeculae was conducted on the plane of the section divided into conventional regions and sub-regions consistent with the previously identified trabecular families that provide regions of relatively homogeneous microstructure. Mean trabecular width and percent bone area were calculated at such sites. Our findings indicate that each of mean trabecular width and percent bone area vary within each proximal femur independently from each other, with dependence on site. Both trabecular parameters show significant differences between pairs of sites. We speculate that a high FRAX index at the hip corresponds to a reduced percent bone area among sites that gives a more homogeneous and less site-specific quality to the proximal femur. This phenomenon may render the local tissue less able to carry out the expected mechanical function.     

The effect of the density-modulus relationship selected to apply material properties in a finite element model of long bone

Material property assignment is a critical step in developing subject-specific finite element models of bone. Inhomogeneous material properties are often applied using an equation relating density and elastic modulus, with the density information coming from CT scans of the bone. Very few previous studies have investigated which density-elastic modulus relationships from the literature are most suitable for application in long bone. No such studies have been completed for the ulna. The purpose of this study was to investigate six such density-modulus relationships and compare the results to experimental strains from eight cadaveric ulnae. Subject-specific finite element models were developed for each bone using micro-CT scans. Six density-modulus equations were trialed in each bone, resulting in a total of 48 models. Data from a previously completed experimental study in which each bone was instrumented with twelve strain gauges were used for comparison. Although the relationship that best matched experimental strains was somewhat specimen and location dependent, there were two relations which consistently matched the experimental strains most closely. One of these under-estimated and one over-estimated the experimental strain values, by averages of 15% and 31%, respectively. The results of this study suggest that the ideal relationship for the ulna may lie somewhere in between these two relations.     

Patterns of strain in the macaque tibia during functional activity.

The strain environment of the tibial midshaft of two female macaques was evaluated through in vivo bone strain experiments using three rosette gauges around the circumference of the bones. Strains were collected for a total of 123 walking and galloping steps as well as several climbing cycles. Principal strains and the angle of the maximum (tensile) principal strain with the long axis of the bone were calculated for each gauge site. In addition, the normal strain distribution throughout the cross section was determined from the longitudinal normal strains (strains in the direction of the long axis of the bone) at each of the three gauge sites, and at the corresponding cross-sectional geometry of the bone. This strain distribution was compared with the cross-sectional properties (area moments) of the midshaft. For both animals, the predominant loading regime was found to be bending about an oblique axis running from anterolateral to posteromedial. The anterior and part of the medial cortex are in tension; the posterior and part of the lateral cortex are in compression. The axis of bending does not coincide with the maximum principal axis of the cross section, which runs mediolaterally. The bones are not especially buttressed in the plane of bending, but offer the greatest strength anteroposteriorly. The cross-sectional geometry therefore does not minimize strain or bone tissue. Peak tibial strains are slightly higher than the peak ulnar strains reported earlier for the same animals (Demes et al. [1998] Am J Phys Anthropol 106:87-100). Peak strains for both the tibia and the ulna are moderate in comparison to strains recorded during walking and galloping activities in nonprimate mammals.     

Effect of weather factors on populations of Helicoverpa armigera moths at cotton-based agro-ecological sites

Pheromone trapping was used to monitor populations of the moth Helicoverpa armigera at five cotton‐based agro‐ecological sites – river, vegetable, orchard, forest and clean cultivation (areas under only cotton cultivation) – in the Bahawalpur district, Pakistan. Three locations at each site were chosen and three pheromone traps at each location were installed in cotton fields. Moth catches were recorded at 15–20 day intervals from 24 October 2004 to 19 December 2006. In 2004, the river sites showed the maximum trapped population of H. armigera (0.22/trap) followed by 0.165 per trap at the vegetable sites. Orchard, clean cultivation and forest sites had zero moth catches. In 2005, the river sites again showed the highest trapped population (0.57/trap), followed by clean cultivation (0.45/trap), vegetable (0.44/trap), orchard (0.40/trap) and forest (0.29/trap). The moths appeared during July to December and March to May. In 2006, sites showed non‐significant difference, with a population range of 0.47 to 0.97 moths per trap. On average, river sites peaked at 0.49 per trap, followed by vegetable (0.38), clean cultivation (0.47), orchard (0.35) and forest (0.25) sites. The peak was observed on 3 April 2006, and moths appeared during February to July and October to December. The minimum temperature in river, forest and clean cultivation sites; the maximum temperature in orchard sites; and the average temperature in river, orchard, forest and clean cultivation sites showed significant positive correlations with trapped moth populations. Relative humidity showed significant negative correlation with population at the orchard sites in 2005. All weather factors during 2004 and 2006 showed non‐significant correlations with the moth populations. No model was found to be best fit by multiple linear regression analysis; however, relative humidity, minimum temperature, maximum temperature, minimum temperature and maximum temperature contributed 8.40, 10.23, 2.43, 4.53 and 2.53% to the population fluctuation of the moth at river, vegetable, orchard, forest and clean cultivation sites, respectively.     

Relationship of the body mass index with skinfolds,girths, and bone breadths in canadian men and women aged 20–70 years

The relationships of Quetelet index (w/h²), or body mass index (BMI), with the sum of skinfolds at five sites, two skinfold-corrected limb girths, and two bone breadths were studied in a cross-sectional sample of 12,282 men and 6,593 women aged 20–70 years. The correlations of the BMI with skinfolds (0.50), bone breadths (0.51), and girths (0.58) were too low for individual prediction. Contingency tables of the BMI and sum of skinfold categories further indict its use for the purpose of assessing adiposity status or monitoring change in individuals.     

Bone growth and calcium balance during simulated weightlessness in the rat

Rats, age 28 days, experiencing tail suspension in modified metabolic cages for 1, 2, and 3 wk were compared with littermate controls. Food and water consumption, urinary and fecal Ca excretion, and serum Ca were measured; hearts, fore- and hindlimb bones, skulls, and mandibles were removed for determination of wet, dry, and ash weights and Ca concentration and for histological examination. Weight gain and Ca intake and excretion were the same for both groups; both displayed net Ca gain. Suspended rats had significantly lower wet, dry, and ash weights of femora and tibiae. Dry weights of the humeri and radii/ulnae were moderately higher, and the skull and mandible dry and ash weights were significantly higher in suspended than in control rats. Cortical thickness of the femur, but not humerus, was less in suspended rats. The data are consistent with the hypothesis that bone growth is influenced by the cardiovascular changes associated with tail suspension.     

Invited Review: Pathogenesis of osteoporosis.

Patients with fragility fractures may have abnormalities in bone structural and material properties such as larger or smaller bone size, fewer and thinner trabeculae, thinned and porous cortices, and tissue mineral content that is either too high or too low. Bone models and remodels throughout life; however, with advancing age, less bone is replaced than was resorbed within each remodeling site. Estrogen deficiency at menopause increases remodeling intensity: a greater proportion of bone is remodeled on its endosteal (inner) surface, and within each of the many sites even more bone is lost as more bone is resorbed while less is replaced, accelerating architectural decay. In men, there is no midlife increase in remodeling. Bone loss within each remodeling site proceeds by reduced bone formation, producing trabecular and cortical thinning. Hypogonadism in 20-30% of elderly men contributes to bone loss. In both sexes, calcium malabsorption and secondary hyperparathyroidism increase remodeling: more bone is removed from an ever-diminishing bone mass. As bone is removed from the endosteal envelope, concurrent bone formation on the periosteal (outer) bone surface during aging partly offsets bone loss and increases bone's cross-sectional area. Periosteal apposition is less in women than in men; therefore, women have more net bone loss because they gain less on the periosteal surface, not because they resorb more on the endosteal surface. More women than men experience fractures because their smaller skeleton incurs greater architectural damage and adapts less by periosteal apposition.     

Laurel forests in Tenerife, Canary Islands

The general wood structure, vessel size and distribution along the stem xylem radius and in petioles were studied in Laurus azorica trees living in a Tenerife laurel forest. The fractions of volume occupied by dry matter, water and air in percentage of wood fresh volume were also studied. The wood showed a diffuse-porous structure, with solitary vessels or vessels somewhat clustered in small radially oriented groups. Vessels had a diameter ranging from 20 to 130 µm. This diameter was minimal close to the pith, increased more than 2-fold with age, and reached its maximum width close to the cambium. Vessel density decreased from 36 vessels mm^-2 near the pith to about 13 vessels mm^-2 near the cambium. Accordingly, the lumen area was small in young xylem close to the pith (0.0015 mm²), reaching a value 5 times larger (0.007 mm²) near the cambium than in the centre of the stem. Lumen area of vessels in petioles was about 1.5% of petiole cross-sectional area and thus much lower than in stems. Mean hydraulic diameter of these vessels was about 20 µm, and mean vessel density about 136 per petiole. There were only small differences in proportions of dry matter, water and air along stem radius. The relevance of each one of these fractions in the wood is discussed as evidence of the possible existence of a number of embolized vessels dispersed in the total functional cross-sectional area of the xylem.     

Comparison of strain measurement in the mouse forearm using subject-specific finite element models,strain gaging,and digital image correlation

Mechanical loading in bone leads to the activation of bone-forming pathways that are most likely associated with a minimum strain threshold being experienced by the osteocyte. To investigate the correlation between cellular response and mechanical stimuli, researchers must develop accurate ways to measure/compute strain both externally on the bone surface and internally at the osteocyte level. This study investigates the use of finite element (FE) models to compute bone surface strains on the mouse forearm. Strains from three FE models were compared to data collected experimentally through strain gaging and digital image correlation (DIC). Each FE model was assigned subject-specific bone properties and consisted of one-dimensional springs representing the interosseous membrane. After three-point bending was performed on the ulnae and radii, moment of inertia was determined from microCT analysis of the bone region between the supports and then used along with standard beam analyses to calculate the Young’s modulus. Non-contact strain measurements from DIC were determined to be more suitable for validating numerical results than experimental data obtained through conventional strain gaging. When comparing strain responses in the three ulnae, we observed a 3–14% difference between numerical and DIC strains while the strain gage values were 37–56% lower than numerical values. This study demonstrates a computational approach for capturing bone surface strains in the mouse forearm. Ultimately, strains from these macroscale models can be used as inputs for microscale and nanoscale FE models designed to analyze strains directly in the osteocyte lacunae.     

Intra-annual tree-ring parameters indicating differences in drought stress of Pinus sylvestris forests within the Erico-Pinion in the Valais (Switzerland)

The effects of drought on radial growth of Pinussylvestris were investigated by comparing sites along hydricgradients. The gradients were located in Valais, an inner Alpine dry valley inSwitzerland, with each consisting of two site types, an extreme dry, xeric siteand a less dry, moderate site. The two site types were assigned tophytosociological associations within the Erico-Pinion. The investigationcovered the responses of tree growth to climate and particularly concentratedonintra-annual features of tree-rings such as earlywood/latewood ratio,intra-annual density fluctuations (IADFs) and traumatic tissues (TTs) as wellasthe sapwood/heartwood ratio. Radial growth differed according to the sitetypes,with trees on dry sites generally showing more missing rings, lower mean ringwidths, lower autocorrelation, higher mean sensitivities, reduced latewoodproportions and lower sapwood areas than trees on moderate sites. Therelationships between climate and tree-ring width, studied using responsefunction analysis, varied strongly between the site types within theErico-Pinion: Tree growth on dry sites was positively influenced byprecipitation at the end of the winter and the beginning of the growing seasonand negatively influenced by temperature in June. Winter precipitation waspositively correlated with radial growth, demonstrating its importance for thesuccessful root and shoot growth of the plants in spring on dry sites. Onmoderate sites, tree growth was less controlled by climate than by priorgrowth.The intra-annual density fluctuations (IADFs) provided a valuable means todifferentiate between the site types. In comparison to the moderate sites, thetrees on dry sites contained more IADFs, and their frequency was increased.Moist-cool conditions in the middle of the growing season were the triggeringfactor for IADFs on dry sites, whereas on moderate sites, there must be anadditional warm period in early summer in order to initiate IADFs. Most IADFswere found in latewood. We found no relationship between climate and traumatictissues (TTs). It is unclear whether other abiotic or biotic factors such aswounding by insects or birds are responsible for the development of TTs. Theassignment of these differences in tree growth behaviour to phytosociologicalassociations will enable a deeper understanding of the site types and willfacilitate the comparison with similar studies. Furthermore, the results can becombined with studies from other scientific disciplines concerning thesephytosociological associations. The ecological indicator values of thevegetation was a precise method for the distinction of site types.     

Mechanical properties of the lateral collateral ligament: effect of cruciate instability in the rabbit

Concomitant soft tissue injury resulting from knee instability following cruciate rupture is a serious clinical problem. To study this injury mechanism, the biomechanical properties of the lateral collateral ligament were measured at 0, 4, 8, 12, and 16 weeks post-operatively in rabbits having the anterior and posterior cruciate ligaments sectioned. No significant changes were found in the ligament's cross-sectional area, tensile mechanical response, or in its hexosamine content. The predominant mode of ligament failure was by bone avulsion at the insertion sites (78 percent) with 86 percent of paired limbs failing in a similar manner.     

A morphometric study of the cochlea of the little brown bat (Myotis lucifugus).

A detailed morphometric study of the basilar membrane was made from serial sections and graphic reconstructions of the cochlea of three little brown bats. Four distinct morphometric changes were observed within the basilar membrane. First, between 0-1.4 mm from the basal end of the cochlea, there is a rapid increase in width and cross-sectional area of the basilar membrane. Secondly, between 1.4-2.5 mm, there is little change in width of the basilar membrane (its cross-sectional area is at its greatest in this region). Thirdly, between 2.7-3.1 mm, there is a sudden decrease in cross-sectional area concomitant with an increase in the width of the basilar membrane. Finally, between 3.1 mm and the apex, there is a gradual decrease in cross-sectional area concomitant with an increase in the width of the basilar membrane. The magnitudes of the cross-sectional areas of the scalae media and vestibuli decrease from base to apex, but this is not true for the scala tympani. The cross-sectional area of the scala tympani appears to decrease from the base to 0.7 mm, then it increases up to 1.4 mm, and then it decreases to the apex. These morphometric changes in the basilar membrane of the little brown bat are compared to those in other echolocating and non-echolocating mammals. The significance of these changes is discussed in relation to the range of hearing in the little brown bat.     

A Comparison of the Soil Water,Nutrient Status,and Litterfall Characteristics of Tropical Heath and Mixed‐Dipterocarp Forest Sites in Brunei1

Two of the main hypotheses to explain the distribution and special characteristics of tropical heath forest are nutrient and water limitation. A study was undertaken to investigate both factors on two sites under tropical heath forest (Badas Forest Reserve) and mixed‐dipterocarp forest (Andulau Forest Reserve) in Brunei. Soil water potentials were monitored at depths of 20, 50, and 90 cm over wet and dry periods for five months at each site. The results showed the mixed‐dipterocarp forest site to be drier at 50 cm depth compared to the tropical heath forest site. There was no significant difference in water potentials between sites at 20 or 90 cm. Nutrient concentrations in the soil solution were monitored at the same depths over a seven‐month period at the same sites. A 12‐month litterfall study was also undertaken to monitor nutrient returns from the canopy at each site. The results of both studies suggest that the tropical heath forest site is poorer in nitrogen, but richer in calcium, than the mixed‐dipterocarp forest site. The results for phosphorus are less clear, but do not suggest that its limitation is a significant factor at the tropical heath forest site compared to the mixed‐dipterocarp forest site. Phosphorus and magnesium concentrations in the soil solution showed a strong positive correlation with sliding 30‐day rainfall totals at both sites.