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ORIGINAL COMMUNICATION The Developing Juvenile Ischium: Macro-Radiographic Insights STEPHEN J. MACLEAN, SUE M. BLACK, AND CRAIG A CUNNINGHAM* Centre for Anatomy and Human Identification, College of Arts, Science and Engineering, University of Dundee, Dundee DD1 5EH Despite the importance of the human pelvis as a weight-bearing structure, there is a paucity of literature that discusses the development of the juvenile innominate from a biomechanical perspective. This study aims to add to the limited body of literature pertaining to this topic through the qualitative analy- sis of the gross architecture of the human ischium during the juvenile period. Macro-radiographs of 55 human ischia ranging from 28 intra-uterine weeks to 14 years of age were examined using intensity-gradient color mapping to high- light changes in gross structural morphology with increasing age. A clear pat- tern of maturation was observed in the juvenile ischium with increasing age. The acetabular component and ramus of the ischium consistently displayed low bone intensity in the postnatal skeletal material. Conversely the posterior body of the ischium, and in particular the ischial spine and lesser sciatic notch, exhibited increasing bone intensity which first arose at 1–2 years of age and became more expansive in older cohorts. The intensity patterns observed within the developing juvenile ischium are indicative of the potential factors influencing the maturation of this skeletal element. While the low intensity ace- tabular fossa indicates a lack of significant biomechanical interactions, the pos- terior increase in bone intensity may be related to the load-bearing nature of the posterior ischium. Clin. Anat. 27:906–914, 2014. VC 2014 Wiley Periodicals, Inc. Key words: ischium; juvenile; macro-radiography; growth; biomechanics INTRODUCTION Over the last two centuries, the internal structure of human bone has been studied extensively in an effort to better understand the role of trabecular bone, its impli- cations for biomechanical competency and the factors which influence the development of its micro- architecture (von Meyer, 1867; Wolff, 1870, 1892; Sato et al., 1986; Tobin, 1955; Huiskes, 2000; Huiskes et al., 2000; Nuzzo et al., 2003; Agarwal et al., 2004; Macho et al., 2005; Cunningham and Black, 2010). A substan- tial number of these investigations have focused on a relatively limited range of skeletal elements, primarily the major long bones (von Meyer, 1867; Salle et al., 2002; Ryan and Krovitz, 2006; Perilli et al., 2008; Gos- man and Ketcham, 2009; Kraus et al., 2009; Barvencik et al., 2010; Hammer, 2010; Ryan and Walker, 2010; Liu et al., 2011; Saparin et al., 2011; Reissis and Abel, 2012). More recently, there has been an expansion into regions of the skeleton previously neglected in terms of trabecular bone analysis, including weight-bearing ele- ments such as the ilium (Sato et al., 1986; Volpato et al., 2008; Cunningham and Black, 2009a,b, 2010; Abel and Macho, 2011), vertebral bodies (Nuzzo et al., 2003; Rapillard et al., 2006), the calcaneus (Maga et al., 2006; Rupprecht et al., 2006) and the talus (Pal and Routal, 1998). However, in spite of this expanding literature base, little is known about the biomechanical loading and resultant cortical and trabecular bone dynamics of many skeletal elements, including the pel- vic girdle (Dalstra et al., 1993). *Correspondence to: Dr. Craig A Cunningham, Centre for Anatomy and Human Identification, University of Dundee, Dow Street, Dundee DD1 5EH. E-mail: c.a.cunningham@dundee.ac.uk Received 18 February 2014; Accepted 18 February 2014 Published online 17 March 2014 in Wiley Online Library (wileyonlinelibrary.com). DOI: 10.1002/ca.22391 VVC 2014 Wiley Periodicals, Inc. Clinical Anatomy 27:906–914 (2014) The pelvic girdle represents a key junctional com- plex for the transfer of weight from the head, trunk and upper extremities to the lower limb (Cunningham and Black, 2009a; Tardieu et al., 2013), therefore it should be an important focus for biomechanical and osteological research. However, the extant literature which describes pelvic bone structure is primarily restricted to phylogenetic studies concentrating on the differences between Homo sapiens and other primate species (Macchiarelli et al., 1999; Martin�on-Torres, 2003; Volpato et al., 2008), with emphasis typically placed on iliac shape and structure. These studies focus almost exclusively on the trabecular patterns of the adult ilium and typically neglect to consider juve- nile bone development. Some papers have however examined the evolutionary importance of variations in parameters such as the ratio between ischial and iliac length (Williams and Orban, 2007) and also the evolu- tion of the morphology of the ischial spine (Abitbol, 1988). As the internal structure of an element is inherently linked to its mechanical environment (Huiskes et al., 2000), a detailed understanding of the developing architecture may provide insight into the driving processes behind each stage of skeletal maturation. The structure of the adult human innominate has been described as a ‘sandwich construction’ consisting of a core of relatively isotropic trabecular bone con- tained between two thin cortical shells (Dalstra et al., 1993). Discussions of discrete trabecular bundles within the ischium specifically document a single tra- jectory: the sacro-ischial trabecular bundle (Aiello and Dean, 1990). This bundle is described as passing between the auricular surface of the ilium and the ischial tuberosity, with the proposed function of trans- mitting upper body weight through the pelvic complex when in a seated position (Scheuer and Black, 2000). This bundle was also described as inter-connecting with the trabeculae of the acetabular margin, perhaps to assist in the distribution of forces associated with the acetabulum during bipedal locomotion (Kapandji, 2011). In a finite element (FE) analysis of the adult innominate, patterns of stress distribution were pro- posed, with consideration given to the load transferred to the femoral head at the acetabulum in conjunction with the action of 21 muscles that insert on the pelvis (Dalstra and Huiskes, 1995). It was concluded that both the cortical bone in the superior portion of the body of the ischium and the acetabular component experienced high levels of stress during different phases of the walking cycle, while limited forces were experienced by the trabecular element of the bone. However, it has been noted subsequently that there were discrepancies in the authors’ consideration of the trabecular bone (Cunningham and Black, 2009a) and therefore caution should be used when interpreting the trabecular models produced. Beyond this FE analysis, few studies have docu- mented the biomechanical environment of the ischium, nor described the factors that may influence the development of this element. However, an understanding of the specific soft- tissue interactions and potential biomechanical influ- ences acting on the bone at each stage of develop- ment can be used to inform initial hypotheses regarding the drivers of skeletal growth, based on an appreciation of gross structural changes through development. This study forms the first stage of an on-going investigation that will examine the developing struc- ture of the juvenile human ischium, with particular reference to the changing biomechanical environment of the pelvis during growth and development. This preliminary study documents qualitatively the gross patterns in bone development within the ischium dur- ing the juvenile period to provide insights into poten- tial driving factors behind the maturation of the ischium, and also to highlight regions of importance for future quantitative analysis. MATERIALS AND METHODS Fifty-five ischia were available to study from the Scheuer Collection of juvenile skeletal remains held within the Centre for Anatomy and Human Identifica- tion, University of Dundee). This unique collectionis an active osteological repository for juvenile human skeletal remains and comprises over 120 subadult individuals from a variety of archaeological, anatomi- cal and forensic sources. The specimens included within this study ranged from 28 intra-uterine (i.u.) weeks to 14 years of age and each specimen was assigned to one of seven age groups based on the estimated age held on record for the collection (Table 1). Only those individuals who did not exhibit evi- dence of ilio-ischial fusion were selected included for analysis. Radiography Each specimen was subjected to macro- radiography using a Multix Tube and Table (Siemens). Specimens were placed with the internal (pelvic) sur- face towards the radiographic plate and the external (acetabular) surface towards the x-ray source. The exposure factors applied were 47 kV, 2 mA, Fine Focus, with an Agfa film screen combination. A focus film distance of 140 cm and an object film distance of 30 cm were used. Macro-radiography is used to mag- nify a radiographic image relative to the size of the object: this is achieved by increasing the object film TABLE 1. Age Groups, Specimen Numbers, and Element Distribution in This Study Age Individuals Left ischia Right ischia Total elements (n) 28 intra-uterine weeks 1 1 1 2 Neonatal 3 3 3 6 426 months 3 3 3 6 1–2 years 3 2 3 5 3–5 years 6 6 5 11 6–9 years 6 6 5 11 10–14 years 10 7 7 14 Total 32 28 27 55 The Developing Juvenile Ischium 907 distance relative to a fixed focal film distance (David- son and Bowman, 2002). Macro-radiography was established as a useful technique in the gross analysis of patterns in trabecular architecture when applied to the developing ilium (Cunningham and Black, 2009a). Color-Mapping and Gradient Analysis To create color gradient maps of the macro- radiographs, each radiograph was converted into a digital image using a flatbed scanner with a superiorly mounted radiographic light. The scan resolution was set to 4,800 dpi to maximize data capture during the digitization process. Each radiograph was scanned as a gray-scale image comprising a standard 256 levels of gray, saved as a .JPEG file without compression. The digital radiographs were opened in Photoshop CS5 (Adobe Systems Inc.), which was used to color the radiographs based on the levels of gray present using the gradient map tool (Fig. 1). To achieve this, the 256 levels of gray were divided into four discreet groups, each of which was assigned a single contrasting color. As the original radiographs were not captured digitally, slight variations in pixel intensities were present in the final JPEG images. This required that each specimen be mapped individually based on the specific grey levels present by assigning a base-line color to the intensity associated with the background of the radiograph. The remaining three colors were assigned an equal propor- tion of the remaining grey levels (e.g., where the base- line color was set at 16%, the remaining gray levels would be distributed evenly between the three remain- ing colors at intervals of 21%). Several of the younger specimens (including speci- mens from the fetal, perinatal, 0–6 months, and 1–2 years groups) displayed a narrower range of intensity levels due to the small size of the elements. Smaller, proportional, changes in intensity were visualized in these specimens using “sensitive” gradient maps. To create these sensitive gradient maps a maximum intensity threshold was set at �85% opacity, in addi- tion to the lower background level threshold estab- lished previously. The intermediary grey levels were then distributed evenly between the remaining colors as before. This narrower threshold window did not negatively impact the analyses as the gross patterns evaluated are based on relative differences in local pixel intensity, not absolute variations. The gradient mapping process divided the ischium into a color map comprised of four colors representing different relative grayscale intensities in the image from yellow, which represents the background level of exposure, through violet and orange to blue, which represents the most radiopaque regions i.e., the whiter regions in the original radiographs (Table 2). RESULTS A progressive pattern of gradient map coloration was observed between each of the age groups (Fig. 2). The features highlighted on the specimens presented Fig. 1. Macro-radiograph [a] and gradient map [b] of the ischium (3–5 years). Four distinct grades of density are identified within the ischium, represented by the four colors (yellow, violet, orange, and blue). [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] TABLE 2. Density Gradient Levels and Their Corre- sponding Colors and Opacity Ranges Relative density level Colour Approximate opacity (%) Very Low Yellow 0–30 Low Violet 31–50 Medium Orange 51–70 High Blue 71–100 908 Maclean et al. http://wileyonlinelibrary.com Fig. 2. Gradient maps of specimens from each age group analyzed. (a) 28 intra- uterine weeks; (b) neonatal; (c) 4–6 months; (d) 1–2 years; (e) 3–5 years; (f) 6–9 years; (g) 10–14 years. Areas of interest referred to in the text are identified by num- ber (1–7). Images are not shown to scale. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] The Developing Juvenile Ischium 909 http://wileyonlinelibrary.com in Figure 2 are representative of all specimens within their respective groups. 28 Intra-Uterine Weeks (n52) The majority of the fetal ischium was represented by a relatively uniform medium intensity of bone as illus- trated by a preponderance of orange coloration (Fig. 2a). An area of higher intensity was noted passing antero-inferiorly across the body of the ischium (between the dashed lines in Fig. 2a) that corresponds to a small raised crest visible on the external surface of the bone. Regions of lower intensity were noted in the acetabular fossa (Label 1 in Fig. 2a), the anterior body of the ischium (2) and around the perimeter of the bone. Neonatal (n56) The neonatal cohort demonstrated a similar pattern to the 28 i.u. weeks group, with an expanded region of lower intensity present anterosuperiorly defining the region of the acetabular fossa (Label 1 in Fig. 2b). A lower intensity region was also identified posterior to the immature acetabular margin (3) in the approxi- mate location of the dominant nutrient foramen. The anterior body of the ischium in this group exhibited a more uniform, higher intensity than in the 28 i.u. weeks cohort but the periphery of the bone retained a lower intensity violet coloration. 4–6 Months (n56) Several observations from this age group were con- sistent with the younger groups: lower intensity regions were evident both anterosuperiorly in the region of the acetabular fossa (1) and inferior to the acetabular margin (3) (Fig. 2c). A lower intensity region was also observed in the anterior body of the ischium (2) distinct from the orange mid-level inten- sity of the posterior body (separated by the dashed line). The developing ramus was also represented by a region of lower intensity bone (4). It was also noted that in two specimens in this group, Regions 1, 3, and 4 are of such low relative intensity that they approxi- mate the background intensity level, with yellow areas visible which were not apparent in either of the younger age cohorts. 1–2 Years (n55) The specimens within this age cohort exhibited sev- eral features consistent with previous age groups (Fig. 2d). Very low intensity regions were evident in the ace- tabular fossa (1), anterior body of the ischium (2), and ramus (4), each with a preponderance of yellow indicat- ing an intensity level very close to background density levels. The difference in relative intensity between the anterior and posterior regions of the body of the ischium was readily apparent in this group (separated by the dashed line). The most significant observation in this group was the developmentof a higher intensity region on the posterior border of the ischium (5), in the region of the lesser sciatic notch. A region of lower intensity bone was also identified along the postero-inferior bor- der of the ischium in the region of the metaphyseal sur- face of the future ischial tuberosity (6). 3–5 Years (n511) Several observed regions in the 3–5 years group were consistent with the patterns described in younger age cohorts (Fig. 2e). The acetabular fossa (1) was defined by a region of low intensity, while the ramus (4) was represented by a region of low intensity proximally which transitioned to a lower intensity at its extremity. The anterior–posterior disparity (separated by the dashed line) was also present in this group, although it was noted that while the anterior body of the ischium maintained a relatively low intensity (2) it did not exhibit intensity levels approximating the background level as previously observed. The higher intensity region in the posterior border of the ischial body around the lesser sciatic notch (5) was also present in this group, and was more extensive than observed in the 1– 2 years cohort. 6–9 Years (n511) This age cohort contained many of the features iden- tified in previous age groups, including lower intensity regions in the acetabular fossa (1) and the ramus (4). The higher intensity region observed in the posterior body of the ischium was also present in the age group (5), surrounded by an area of medium intensity (orange) (Fig. 2f). The differentiation between the lower intensity anterior body (2) and higher intensity posterior body of the ischium was also well defined in this age group (separated by dashed line). 10–14 Years (n514) The 10–14 years age group exhibited several inten- sity patterns that were consistent with observations of the younger age groups, including the very low intensity regions associated with the acetabular fossa (1) and the ramus (4) (Fig. 2g). In four specimens, the ramus of the ischium had fused to the pubic bone; however, there were no obvious differences evident in the intensity of the ischiopubic rami of the fused and unfused speci- mens. The intensity gradient between the anterior and posterior body of the ischiumwas also identifiable in this age cohort (separated by the dashed line), with a lower intensity area evident in the anterior body of the ischium (2). However, the distinction between these two regions was less pronounced than previously observed. A more expansive higher intensity region around the lesser sci- atic notch was observed in this cohort (5). Seven out of ten specimens also exhibited variable regions of mid to high density bone curving posteriorly in the approxi- mate location of the anterior extremity of the develop- ing acetabularmargin (7). DISCUSSION The radiographic representations of the specimens in this study demonstrate a largely progressive 910 Maclean et al. pattern of intensity distribution and development with increasing maturity. The ischium transitioned from a near-uniform, undifferentiated distribution of bone in the fetal period to adopt a more regionalized pattern with defined variations in intensity in the early adoles- cent period. A recognizable mature morphology began to manifest in the 1–2 year age group and was clearly established by the age of 3–5 years. It is important to consider the limitations of any imaging modality when interpreting results, particu- larly with regards to radiographic imaging technolo- gies. The most significant limitation of plane plate radiography is the superimposition of structures on a single plane, which is a result of the compression of a three dimensional structure into a two dimensional image (Hsieh, 2003). As a consequence of this com- pression, variations in the three-dimensional structure along the z-axis (in the plane of the radiographic beam) will affect beam attenuation and create varia- tions in the pixel intensities of the radiograph. To mini- mize the confuscating effect of superimposition, each gradient map was considered in conjunction with the specimen from which the radiograph was obtained. This made it possible to identify variations in intensity that could not be attributed solely to the development of gross external structures. It must also be noted that radiography does not allow a distinction of tra- becular and cortical bone tissues; rather it creates an image which is a composition of both types of bone tissue. All variations in intensity therefore represent “combined bone intensity” and highlight relative dif- ferences in structure within an element and not abso- lute differences between specimens. Because of the limited number of specimens avail- able, the youngest group in this study consisted of a single individual of 28 i.u. weeks, including both the left and right ischium. The small group size, combined with the small size of these specimens and the rela- tively low density of bone formed, precludes definitive assessment of the prenatal structure of the ischium. Rather, this group was included to provide an indica- tion of the potential structure of the ischium at this age; however, the results should be treated with cau- tion. Similarities were observed between these speci- mens and the more mature, postnatal skeletal material, for example the lower intensity region in the acetabular fossa. However, these variations in radiolu- cency may also be attributed to gross variations in external morphology. For example, the fetal acetabu- lar fossa was represented by a lower, flattened region on the anterosuperior surface while the noted region of increased intensity corresponds to a small raised crest passing antero-inferiorly across the body of the ischium. This relatively nondescript radiographic appearance of the fetal ischium contrasts with observations of the fetal ilium, which displays a precocious and progres- sive pattern of development from the early fetal period (Cunningham and Black, 2009a). Rudimentary buttressing was identifiable in anterior and posterior trajectories consistent with the mature adult iliac structure in specimens as young as 23–30 intra- uterine weeks. In adult specimens, these trajectories are purported to be responsible for the transfer of weight between the auricular surface of the ilium and the femoral head at the acetabulum (Aiello and Dean, 1990; Scheuer and Black, 2000; Kapandji, 2011), and also to assist in the distribution of associated tensile forces (Macchiarelli et al., 1999). The discovery of cor- responding, although immature, trajectories in the fetal ilium raised questions as to the cause of such precocious development, as the pelvis is not a weight- bearing structure prior to birth (Walker, 1991) and tra- ditional models postulate that bone structure is pri- marily dictated by mechanical stimuli (Wolf, 1892; Huiskes, 2000). It was proposed that the early estab- lishment of supportive trajectories may also be influ- enced by genetic and non-load-bearing mechanical interactions (Cunningham and Black, 2009a,b), although subsequent publications also stressed the importance of vascular invasion as a factor which may direct trabecular growth in the ilium (Cunningham and Black, 2010). The absence of such precocious devel- opment in the fetal ischium included in this study may be indicative of a different set of osteogenic instiga- tors for the ischium, although it is not possible to sug- gest alternative influences from the study of a single individual. Analysis of the neonatal and postnatal development of the ischium revealed progressive patterns of den- sity in several key regions arising after birth. In both the neonatal and 4–6 months age groups, a small elliptical region was identified inferior to the acetabu- lar rim that was characterized by a lower bone density relative to the surrounding bone. It is proposed that this lower region of density is related to vascular spaces present within the ischium, supported by the presence of a nutrient foramen located in theexternal cortical shell in this region. It has been reported that the acetabular branch of the inferior gluteal artery provides the arterial supply to the ischium (Beck et al., 2003) and the proliferation of blood vessels has been shown to emanate from this approximate region in the neonatal ischium (Crock, 1996). Angiogenesis has previously been demonstrated to be of great importance in bone formation (Brandi and Collin- Osdoby, 2006) and has been suggested to be highly influential in the development of the internal architec- ture of the neonatal ilium (Cunningham and Black, 2010). It is acknowledged that arterial invasion of a cartilaginous template will create vascularized spaces within the internal architecture (Eriksen et al., 2007), which may in turn manifest as an apparent decrease in trabecular bone volume present in this location. Throughout the developmental period investigated in this study, the acetabular fossa was consistently revealed to be amongst the least dense regions of the ischium. While this may be partially attributed to the relatively shallow cross-sectional depth at this point, it may also be evidence of the absence of any significant biomechanical forces acting across this region. The acetabular fossa forms the nonarticular floor of the acetabulum and is therefore unlikely to be involved directly in the transfer of mechanical loading (Daniel et al., 2005): this transfer is instead described as occurring across the lunate surface, which constitutes the articular surface of the acetabulum (Scheuer and Black, 2000). Several authors have indicated that the The Developing Juvenile Ischium 911 principal contact between the lunate surface and fem- oral head when bearing weight occurs in the superior dome of the lunate surface (Greenwald and Haynes, 1972; Ipavec et al., 1999; Daniel et al., 2005; Chuck- paiwong et al., 2009), which is located in the iliac con- tribution to the acetabulum. The forces experienced by the remaining lunate surface are less intense and vary with types and phases of movement (Ipavec et al., 1999); for example, load-bearing contact within the ischial region of the acetabulum is intermittent during walking (Greenwald and Haynes, 1972). This diminished level of force in the ischial component of the acetabulum may explain the discrepancy between the relatively low volume of bone present in this region compared with the densities observed in the ilium (Cunningham and Black, 2009a), and subse- quent quantitative data supporting increased trabecu- lar bone reinforcement of the acetabular roof (Cunningham and Black, 2009b). Throughout postnatal development the posterior body of the ischium maintained a higher intensity than the anterior body, bridging the region between the acetabulum and the metaphyseal surface of the ischial tuberosity. From 1 to 2 years of age, an ellipti- cal, very high intensity region developed adjacent to the developing lesser sciatic notch. This very high intensity region extends with increasing age to bridge the posterior border of the ischium between the ischial tuberosity’s metaphyseal surface and the inferior aspect of the ischial spine. Several potential causative factors can be suggested for these increases in intensity. The region of very high intensity originally forms in the lesser sciatic notch, which contributes to the bony margins of the lesser sci- atic foramen in life. Several structures pass through this foramen, most importantly the internal pudendal artery and the pudendal nerve (Standring, 2008). Evidence from the literature suggests that both vascular (Lauren- son, 1964; Brandi and Collin-Osdoby, 2006; Eriksen et al., 2007; Cunningham and Black, 2010) and neuro- logical tissues (Laurenson, 1964; Garc�ıa-Castellano et al., 2000; Jones et al., 2004; Wang et al., 2006) have an influence on skeletal growth and metabolic regula- tion, although the precise mechanisms by which this influence is exerted are currently unknown (Garc�ıa-Cas- tellano et al., 2000). Therefore, it is possible that this ini- tial increase in intensity is a result of a neurovascular- induced increase in bone deposition in the region of the lesser sciatic notch. As neurovascular structures in the pelvis are estab- lished in the embryonic period (Laurenson, 1964; Sche- uer and Black, 2000; Sadler, 2010) it would be expected that any increases in density caused by the presence of these structures would also be evident in the younger groups. The significance of the age at which the region of increased intensity arises must also be considered. The ages of 1–2 years follows the initial development of unassisted upright standing and walking between 9 and 18 months (WHOMGRSG, 2006). Therefore it is impor- tant to consider the alteration in load transfer and those muscles and ligaments which are involved in a bipedal gait as potential mechanisms driving an increase in bone intensity in this region. The forces passing through the pelvis during walking were not only described as passing through the ilium to the superior acetabulum (Dalstra and Huiskes, 1995), but it was also observed that the ischium transmitted a degree of force through the body and acetabular rim during several of the gait phases. While Dalstra and Huiskes (1995) proposed that this force is primarily transmitted through the corti- cal shell, with only a limited trabecular involvement, specific ischial acetabular trabeculae have been described as radiating posteriorly from the acetabulum to assist in the dispersal of forces (Kapandji, 2011). An additional trabecular trajectory, the sacro-ischial tra- becular bundle, has also been described in the ischium as arising between the auricular surface of the ilium and the ischial tuberosity, to transmit the load of the upper body while in a seated position (Aiello and Dean, 1990; Scheuer and Black, 2000; Kapandji, 2011). The devel- opment of unassisted sitting precedes walking, arising between 4 and 9 months (WHOMGRSG, 2006), which may also be a factor in the observed increase in bone intensity in the posterior border of the ischium from 1 to 2 years of age. The posterior body of the ischium has also been described as an important location for muscular and lig- amentous attachment to support both the upright orien- tation of the pelvis (Scheuer and Black, 2000) and the pelvic viscera (Abitbol, 1988). Insertions on the poste- rior aspect of the ischium include the sacrospinous and sacrotuberous ligaments, the hamstring muscle group and numerous small pelvic muscles (Standring, 2008). The actions of these structures generate varying forces within the posterior ischium; for example the hamstring group have been described as creating a relatively high level of force during the walking cycle (Crowninshield and Brand, 1981). The sacrospinous and sacrotuberous ligaments are described as being important inmaintain- ing the upright position of the pelvis while bearing weight and resisting anterior rotation at the sacro-iliac joint (Scheuer and Black, 2000). It has subsequently been suggested that the sacrotuberous ligament does not have a significant load-bearing role (Woodley and Kennedy, 2005) while the sacro-spinous ligament, in association with the iliococcygeal muscles, have been described as generating high tensile stresses between the sacrum and ischium (Tardieu et al., 2013). However, no data has been published describing the specific forces acting through either of these ligaments, there- fore this postulationmust be consideredwith caution. There is a well-established relationship which dic- tates that mechanical strains trigger the remodelling and optimization of both cortical and trabecular bone structure (Wolff, 1892; Huiskes, 2000; Huiskes et al., 2000; Ruff et al., 2006). Therefore, it is reasonable to expect that the high number of muscular interactions, ligamentous constraints and force transmissions should result in a reinforcement of the skeletal archi- tecture in the region. That the high-density region arises after the attainmentof bipedal locomotion lends further support to this hypothesis. CONCLUSIONS Macro-radiographic analysis of the ischium has revealed several distinctive patterns of bone intensity 912 Maclean et al. arising during the development of this skeletal ele- ment. Evidence of vascular invasion during infancy may be identifiable inferior to the ischial acetabular margin. From the age of 1–2 years, areas of increased intensity develop in the posterior region of the body of the ischium, which are consistent with the reinforce- ment of the skeletal architecture in this location. This may be a response to the biomechanical forces attrib- uted to the development of adult posture and gait in addition to the forces imparted through soft tissue interactions in this region, although potential neuro- genic influences in the lesser sciatic notch should not be discounted. Evidence also indicates that the ischial contribution to the acetabulum may not have a signifi- cant role in the transfer of force to the femoral head, due to the absence of reinforcement to the bone in this region. However, it must be reiterated that this investiga- tion forms only the preliminary stage of a larger inves- tigation to quantify changes to the structure of the developing ischium. ACKNOWLEDGMENTS The authors thank Margaret Low for assistance with radiographic procedures and for sharing her expertise and insight. 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