Mechanical Significance of Femoral Head Trabecular
Bone Structure in Loris and Galago Evaluated Using
Micromechanical Finite Element Models
Timothy M. Ryan
1
* and Bert van Rietbergen
2
1
Department of Anthropology and Center for Quantitative Imaging, The Pennsylvania State University, University
Park, Pennsylvania 16802
2
Department of Biomedical Engineering, Eindhoven University of Technology, 5600 MB Eindhoven, The Netherlands
KEY WORDS finite element analysis; femur; trabecular bone; biomechanics
ABSTRACT Work on the interspecific and intraspe-
cific variation of trabecular bone in the proximal femur of
primates demonstrates important architectural variation
between animals with different locomotor behaviors. This
variation is thought to be related to the processes of bone
adaptation whereby bone structure is optimized to the
mechanical environment. Micromechanical finite element
models were created for the proximal femur of the leaping
Galago senegalensis and the climbing and quadrupedal
Loris tardigradus by converting bone voxels from high-
resolution X-ray computed tomography scans of the fem-
oral head to eight-noded brick elements. The resulting
models had approximately 1.8 million elements each.
Loading conditions representing takeoff phase of a leap
and more generalized load orientations were applied to
the models, and the models were solved using the iterative
“row-by-row” matrix-vector multiplication algorithm. The
principal strain and Von Mises stress results for the leap-
ing model were similar for both species at each load ori-
entation. Similar hip joint reaction forces in the range of
4.9 to 12 body weight were calculated for both species
under each loading condition, but the hip reaction values
estimated for Loris were higher than predicted based on
locomotor behavior. These results suggest that functional
adaptation to hip joint loading may not fully explain the
differences in femoral head trabecular bone structure in
Galago and Loris. The finite element method represents a
unique and useful tool for analyzing the functional adap-
tation of trabecular bone in a diversity of animals and for
reconstructing locomotor behavior in extinct taxa. Am J
Phys Anthropol 126:82–96, 2005. © 2004 Wiley-Liss, Inc.
Recent work on the interspecific variation of tra-
becular bone in the proximal femur of extant pri-
mates has revealed important architectural differ-
ences between taxa with divergent locomotor
behaviors (Fajardo and Mu ¨ ller, 2001; MacLatchy
and Mu ¨ ller, 2002; Ryan, 2001; Ryan and Ketcham,
2002b; but see Viola, 2002). It is generally thought
that, just as with external bone anatomy and diaph-
yseal cross-sectional geometry (Anemone, 1990;
Connour et al., 2000; Demes and Jungers, 1989),
this variation reflects differences in joint loading
and limb use during various activities. It is not
known, however, whether these structural differ-
ences can be fully explained as the result of epige-
netic or evolutionary functional adaptation, or
whether other nonstructural causes such as genetic,
nutritional, hormonal, or gender factors may play a
role as well.
Experimental and modeling work over the past
few years has further developed our understanding
of the mechanisms of bone modeling (Huiskes et al.,
2000; Sommerfeldt and Rubin, 2001). It is now rel-
atively well-established that the local bone modeling
process is governed by mechanical feedback where
osteoblasts add bone to heavily loaded sites, and
osteoclasts remove bone from sites with lower tissue
loadings (Hsieh et al., 2001; Huiskes et al., 2000).
This response is facilitated by the process of mech-
anotransduction, involving the lacuno-canilicular
porosity of the bone matrix (Burger and Klein-Nu-
lend, 1999). A reasonable hypothesis that follows
from these observations, assuming general similar-
ity across mechanosensory cells, is that stresses and
strains at the trabecular bone tissue level should be
distributed rather evenly over the trabecular archi-
tecture within a specific mechanically homogeneous
Grant sponsor: United States National Science Foundation; Grant
number: BCS-9908847; Grant sponsor: Leakey Foundation; Grant
sponsor: Texas Advanced Computing Center, University of Texas at
Austin.
*Correspondence to: Dr. Timothy M. Ryan, Department of Anthro-
pology, 409 Carpenter Building, The Pennsylvania State University,
University Park, PA 16802. E-mail: tmr21@psu.edu
Received 1 May 2003; accepted 9 September 2003.
DOI 10.1002/ajpa.10414
Published online 12 May 2004 in Wiley InterScience (www.
interscience.wiley.com).
AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 126:82–96 (2005)
© 2004 WILEY-LISS, INC.