Predictive estimation of ovine hip joint centers: A regression approach.

Estimation of the hip joint center in ovine biomechanical analysis is often overlooked or estimated using a marker on the greater trochanter which can result in large errors that propagate through subsequent analyses. The purpose of this study was to develop a novel method of estimating the hip joint centers in sheep to facilitate more accurate analysis of ovine biomechanics. CT scans from 16 sheep of varying ages, weight, sex, and phenotypes were acquired and the data was used to calculate the known hip joint center by sphere fitting the femoral head. Anatomical measurements and additional subject information were used to create a variety of regression models to estimate the hip joint centers in absence of CT data. The best regression equation created utilized markers placed on the tuber coxae and tuber ischii of the pelvis and resulted in a mean 3D Euclidean distance error of 6.43 ± 2.22 mm (mean ± standard deviation) between the known and estimated hip joint center. The regression models produced allow for more detailed, accurate and robust analysis of sheep biomechanics.

Comparative life history patterns of female gorillas.

OBJECTIVES: Several theories have been proposed to explain the impact of ecological conditions on differences in life history variables within and between species. Here we compare female life history parameters of one western lowland gorilla population (Gorilla gorilla gorilla) and two mountain gorilla populations (Gorilla beringei beringei). MATERIALS AND METHODS: We compared the age of natal dispersal, age of first birth, interbirth interval, and birth rates using long-term demographic datasets from Mbeli Bai (western gorillas), Bwindi Impenetrable National Park and the Virunga Massif (mountain gorillas). RESULTS: The Mbeli western gorillas had the latest age at first birth, longest interbirth interval, and slowest surviving birth rate compared to the Virunga mountain gorillas. Bwindi mountain gorillas were intermediate in their life history patterns. DISCUSSION: These patterns are consistent with differences in feeding ecology across sites. However, it is not possible to determine the evolutionary mechanisms responsible for these differences, whether a consequence of genetic adaptation to fluctuating food supplies ("ecological risk aversion hypothesis") or phenotypic plasticity in response to the abundance of food ("energy balance hypothesis"). Our results do not seem consistent with the extrinsic mortality risks at each site, but current conditions for mountain gorillas are unlikely to match their evolutionary history. Not all traits fell along the expected fast-slow continuum, which illustrates that they can vary independently from each other ("modularity model"). Thus, the life history traits of each gorilla population may reflect a complex interplay of multiple ecological influences that are operating through both genetic adaptations and phenotypic plasticity.

Late Pleistocene osseous projectile point from the Manis site, Washington-Mastodon hunting in the Pacific Northwest 13,900 years ago.

Bone fragments embedded in a rib of a mastodon (Mammut americanum) from the Manis site, Washington, were digitally excavated and refit to reconstruct an object that is thin and broad, has smooth, shaped faces that converge to sharp lateral edges, and has a plano-convex cross section. These characteristics are consistent with the object being a human-made projectile point. The 13,900-year-old Manis projectile point is morphologically different from later cylindrical osseous points of the 13,000-year-old Clovis complex. The Manis point, which is made of mastodon bone, shows that people predating Clovis made and used osseous weapons to hunt megafauna in the Pacific Northwest during the Bølling-Allerød.

Population dynamics of western gorillas at Mbeli Bai.

Long-term studies of population dynamics can provide insights into life history theory, population ecology, socioecology, conservation biology and wildlife management. Here we examine 25 years of population dynamics of western gorillas at Mbeli Bai, a swampy forest clearing in Nouabalé-Ndoki National Park, the Republic of Congo. The Mbeli population more than doubled from 101 to 226 gorillas during the study. After adjusting for a net influx of gorillas into the study population, the increase represents an inherent growth rate of 0.7% per year, with 95% confidence limits between -0.7% and 2.6%. The influx of gorillas mainly involved immigration of individuals into existing study groups (social dispersal), but it also included the appearance of a few previously unknown groups (locational dispersal). The average group size did not change significantly during the study, which is consistent with the possibility that western gorillas face socioecological constraints on group size, even when the population is increasing. We found no significant evidence of density dependence on female reproductive success or male mating competition. The distribution of gorillas among age/sex categories also did not change significantly, which suggests that the population had a stable age structure. Our results provide evidence of population stability or growth for some western gorillas (albeit within a small area). The results highlight the value of law enforcement, long-term monitoring, and protected areas; but they do not diminish the importance of improving conservation for this critically endangered species.

Dissecting the two mechanisms of scramble competition among the Virunga mountain gorillas.

ABSTRACT: Two mechanisms have been proposed to explain why scramble competition can increase the travel requirements of individuals within larger groups. Firstly, individuals in larger groups may be more likely to encounter food sites where other group members have already eaten, leading to greater asynchronous "individual" travel to find fresh sites. Secondly, when food sites are aggregated into patches, larger groups may need to visit more patches to obtain the same amount of food per capita, leading to greater synchronous "group" travel between patches. If the first mechanism can be mitigated by increasing group spread, then we expect the second mechanism to be more sensitive to group size. Here, we examine the individual travel and group travel of the Virunga mountain gorillas, along with potential implications for the two mechanisms of scramble competition. Asynchronous individual travel accounted for 67% of the total travel time, and the remainder arose from group travel. Group spread increased significantly for larger groups, but not enough to prevent an increase in individual travel. Contrary to expectations, group travel decreased with size among most groups, and we found only limited evidence of patch depletion that would cause the second mechanism of scramble competition. Collectively, our results illustrate how the influence of group size can differ for individual travel versus group travel, just as it differs among species for overall travel. Studies that distinguish between the two mechanisms of scramble competition may enhance our understanding of ecological constraints upon group size, including potential differences between frugivores and folivores. SIGNIFICANCE STATEMENT: Feeding competition provides insight into how group size can influence the foraging patterns of social animals, but two key mechanisms are not typically compared. Firstly, larger groups may visit more patches to access the same amount of food per capita (group travel). Secondly, their individuals may also need to move past more spots where another member has already eaten (individual travel). Contrary to expectations, we found that group travel decreased with size for most groups of mountain gorillas, which may reflect extra travel by smaller groups to avoid larger groups. Individual travel increased with size in most groups, even though gorillas in larger groups compensated by spreading out over a broader area. The two mechanisms revealed patterns that were not apparent in our previous study of overall travel. Our approach may help to explain potential differences between folivores and frugivores.

Methodology for performing biomechanical push-out tests for evaluating the osseointegration of calvarial defect repair in small animal models.

Push-out tests are frequently used to evaluate the bone-implant interfacial strength of orthopedic implants, particularly dental and craniomaxillofacial applications. There currently is no standard method for performing push-out tests on calvarial models, leading to a variety of inconsistent approaches. In this study, fixtures and methods were developed to perform push-out tests in accordance with the following design objectives: (i) the system rigidly fixes the explanted calvarial sample, (ii) it minimizes lateral bending, (iii) it positions the defect accurately, and (iv) it permits verification of the coaxial alignment of the defect with the push-out rod. The fixture and method was first validated by completing push-out experiments on 30 explanted murine cranial caps and two explanted leporine cranial caps, all induced with bilateral sub-critical defects (5.0 mm and 8.0 mm nominal diameter for the murine and leporine models, respectively). Defects were treated with an autograft (i.e., excised tissue flap), a shape memory polymer (SMP) scaffold, or a PEEK implant. Additional validation was performed on 24 murine cranial caps induced with a single, unilateral critically-sized defect (8.0 mm nominal diameter) and treated with an autograft or a SMP scaffold.•A novel fixture was developed for performing push-out mechanical tests to characterize the strength of a bone-implant interface in calvarial defect repair.•The fixture uses a 3D printed vertical clamp with mating alignment component to fix the sample in place without inducing lateral bending and verify coaxial alignment of push-out rod with the defect.•The fixture can be scaled to different calvarial defect geometries as validated with 5.0 mm bilateral and 8.0 mm single diameter murine calvarial defect model and 8.0 mm bilateral leporine calvarial defect model.

Evaluation of a self-fitting, shape memory polymer scaffold in a rabbit calvarial defect model.

Self-fitting scaffolds prepared from biodegradable poly(ε-caprolactone)-diacrylate (PCL-DA) have been developed for the treatment of craniomaxillofacial (CMF) bone defects. As a thermoresponsive shape memory polymer (SMP), with the mere exposure to warm saline, these porous scaffolds achieve a conformal fit in defects. This behavior was expected to be advantageous to osseointegration and thus bone healing. Herein, for an initial assessment of their regenerative potential, a pilot in vivo study was performed using a rabbit calvarial defect model. Exogenous growth factors and cells were excluded from the scaffolds. Key scaffold material properties were confirmed to be maintained following gamma sterilization. To assess scaffold integration and neotissue infiltration along the defect perimeter, non-critically sized (d = 8 mm) bilateral calvarial defects were created in 12 New Zealand white rabbits. Bone formation was assessed at 4 and 16 weeks using histological analysis and micro-CT, comparing defects treated with an SMP scaffold (d = 9 mm x t = 1 or 2 mm) to untreated defects (i.e. defects able to heal without intervention). To further assess osseointegration, push-out tests were performed at 16 weeks and compared to defects treated with poly(ether ether ketone) (PEEK) discs (d = 8.5 mm x t = 2 mm). The results of this study confirmed that the SMP scaffolds were biocompatible and highly conducive to bone formation and ingrowth at the perimeter. Ultimately, this resulted in similar bone volume and surface area versus untreated defects and superior performance in push-out testing versus defects treated with PEEK discs. STATEMENT OF SIGNIFICANCE: Current treatments of craniomaxillofacial (CMF) bone defects include biologic and synthetic grafts but they are limited in their ability to form good contact with adjacent tissue. A regenerative engineering approach using a biologic-free scaffold able to achieve conformal fitting represents a potential "off-the-shelf" surgical product to heal CMF bone defects. Having not yet been evaluated in vivo, this study provided the preliminary assessment of the bone healing potential of self-fitting PCL scaffolds using a rabbit calvarial defect model. The study was designed to assess scaffold biocompatibility as well as bone formation and ingrowth using histology, micro-CT, and biomechanical push-out tests. The favorable results provide a basis to pursue establishing self-fitting scaffolds as a treatment option for CMF defects.

Double Tension Slide Technique as a Novel Repair for Distal Biceps Tendon Tear: A Biomechanical Evaluation.

Background A comparative biomechanical analysis of two distal biceps tendon repair techniques was performed: a single suture tension slide technique (TST) and two suture double tension slide (DTS) technique. Methodology Ten matched pairs of fresh frozen human cadaveric elbows (20 elbows) were randomly separated into two cohorts for distal biceps tendon repair. One cohort underwent the TST, and the other underwent the DTS technique. The tendon was preconditioned with cyclic loading from 0° to 90° at 0.5 Hz for 3,600 cycles with a 50 N load. The specimens were then loaded to failure at a rate of 1 mm/s. The difference in the load to failure between the groups was analyzed using the Student's t test. The mode of failure was compared between groups using the chi-square test. All p-values were reported with significance set at p < 0.05. Results Overall, 77.8% of the included matched pairs demonstrated greater load to failure in the DTS group. The mean load to failure in the DTS group was 383.3 ± 149.3 N compared to 275.8 ± 98.1 N in the TST group (p = 0.13). The DTS specimens failed at the tendon (5/9), suture (3/9), and bone (1/9). The TST specimens failed at the tendon (4/9) and suture (5/9) only. There was no significant difference in failure type between groups (p = 0.76). Conclusions DTS demonstrates a similar to greater load to failure compared to TST with a trend towards statistical significance. The redundancy provided by the second suture has an inherent advantage without compromising the biomechanical testing.

Platelet-rich plasma enhances mechanical strength of strattice in rat model of ventral hernia repair.

Incisional hernia is a common complication of hernia repair despite the development of various synthetic and bio-synthetic repair materials. Poor long-term mechanical strength, leading to high recurrence rates, has limited the use of acellular dermal matrices (ADMs) in ventral hernia repair (VHR). Biologically derived meshes have been an area of increasing interest. Still these materials bring the risk of more aggressive immune response and fibrosis in addition to the mechanical failures suffered by the synthetic materials. Platelet-rich plasma (PRP), a growth-factor-rich autologous blood product, has been shown to improve early neovascularization, tissue deposition, and to decrease the rates of recurrence. Here, we demonstrate that PRP promotes the release of growth factors stromal derived factor (SDF)-1, transforming growth factor-beta, and platelet-derived growth factor in a dose-dependent manner. Additionally, we utilize an aortic ring angiogenesis assay to show that PRP promotes angiogenesis in vitro. A rat model of VHR using StratticeTM ADM demonstrates similar findings in vivo, corresponding with the increased expression of vascular endothelial growth factor and collagen type 1 alpha 1. Finally, we show that the molecular and cellular activity initiated by PRP results in an increased mechanical stiffness of the hernia repair mesh over time. Collectively, these data represent an essential step in demonstrating the utility and the mechanism of platelet-derived plasma in biomaterial-aided wound healing and provide promising preclinical data that suggest such materials may improve surgical outcomes.

Variability of weaning age in mountain gorillas (Gorilla beringei beringei).

OBJECTIVES: Weaning is a key life history milestone for mammals that represents both the end of nutritional investment from the perspective of mothers and the start of complete nutritional independence for the infants. The age at weaning may vary depending on ecological, social, and demographic factors experienced by the mother and infant. Bwindi mountain gorillas live in different environmental conditions and have longer interbirth intervals than their counterparts in the Virunga Volcanoes, yet other life history characteristics of this population remain less well known. We use long-term data from Bwindi Impenetrable National Park, Uganda to examine factors related to weaning age. MATERIALS AND METHODS: We analyzed data on infants born in four mountain gorilla groups in Bwindi to quantify their age of weaning (defined as last nipple contact) and to test if the sex of offspring, parity, and dominance rank of mother influences age of weaning. We also compared the age at weaning and time to conception after resumption of mating in Bwindi and Virunga gorillas. RESULTS: Bwindi gorillas were weaned at an average age of 57.5 months. No difference was found between age of weaning for primiparous and multiparous mothers, nor did maternal dominance rank influence age of weaning, but sons were weaned at a later age than daughters. The majority of Bwindi mothers were still suckling when they resumed mating and mothers generally conceived before they weaned their previous offspring. The age of weaning was significantly later in Bwindi than in Virunga gorillas. After mothers resumed mating, the time to conceiving the next offspring was not significantly longer for Bwindi females than Virungas females (6 vs. 4 months). DISCUSSION: Later weaning age for sons than daughters is similar to findings of other studies of great apes. Bwindi mountain gorillas are weaned at approximately the same age as western gorillas and chimpanzees, which is more than a year later than Virunga mountain gorillas. The results of this study suggest that variation in ecological conditions of populations living in close geographic proximity can result in variation in life history patterns, which has implications for understanding the evolution of the unique life history patterns of humans.

A versatile biaxial testing platform for soft tissues.

Uniaxial testing remains the most common modality of mechanical analysis for biological and other soft materials; however, biaxial testing enables a more comprehensive understanding of these materials' mechanical behavior. In recent years, a number of commercially available biaxial testing systems designed for biological materials have been produced; however, there are common limitations that are often associated with using these systems. For example, the range of allowable sample geometries are relatively constrained, the clamping systems are relatively limited with respect to allowable configurations, the load and displacement ranges are relatively small, and the software and control elements offer relatively limited options. Due to these constraints, there are significant benefits associated with designing custom biaxial testing systems that meet the technical requirements for testing a broad range of materials. Herein we present a design for a biaxial testing system with capabilities that extend beyond those associated with typical commercially available systems. Our design is capable of performing uniaxial tests, traditional biaxial tests, and double lap shear (simple shear) tests, in either a displacement or load control mode. Testing protocols have been developed and proof-of-concept experiments have been performed on commercially available silicone membranes and rat abdominal skin samples.

Addition of platelet-rich plasma supports immune modulation and improved mechanical integrity in Alloderm mesh for ventral hernia repair in a rat model.

The recurrence of ventral hernias continues to be a problem faced by surgeons, in spite of efforts toward implementing novel repair techniques and utilizing different materials to promote healing. Cadaveric acellular dermal matrices (Alloderm) have shown some promise in numerous surgical subspecialties, but these meshes still suffer from subsequent failure and necessitation of re-intervention. Here, it is demonstrated that the addition of platelet rich plasma to Alloderm meshes temporally modulates both the innate and cytotoxic inflammatory responses to the implanted material. This results in decreased inflammatory cytokine production at early time points, decreased matrix metalloproteinase expression, and decreased CD8+ T cell infiltration. Collectively, these immune effects result in a healing phenotype that is free from mesh thinning and characterized by increased material stiffness.

A low-cost, novel endoscopic repeated-access port for small animal research.

Repeated endoscopic access to the abdominal cavity of animal models is useful for a variety of research applications. However, repeated surgical access may affect the welfare of the animal and compromise results. We present the design and benchtop manufacturing process for a self-sealing endoscopic port requiring surgical incision only at implantation. It can be used for repeated body cavity access over a long time period. This device reduces costs, animals required for a given study, and potential suffering for each animal. This novel endoscopic port is designed for low-cost benchtop manufacturing without expensive equipment such as injection molding facilities. Devices manufactured using the method described in this work have been implanted successfully in hen models for investigation of ovarian cancer for over two years. All work followed Texas A&M University institutional guidelines and was covered under Animal Use Protocol 2017-0172, approved by TAMU Animal Care and Use Committee (IACUC). This method can be translated to produce similar devices for use in other small animal models besides the galline model used in this work. This method can be used to produce devices for slightly different purposes than repeated endoscopic access, such as production of an entry port for surgical tools.

Distal Biceps Tendon Repair Using a Double Tension Slide Technique.

Distal biceps tendon ruptures are thought to be secondary to an acute forceful eccentric load on a degenerative tendon. Nonoperative treatment following rupture leads to significantly decreased forearm supination and elbow flexion strength. There are several techniques described in the literature for repair. This article describes, with video illustration, distal biceps tendon repair using a double tension slide technique with 2 No. 2 high-tension nonabsorbable composite sutures.

Clamping soft biologic tissues for uniaxial tensile testing: A brief survey of current methods and development of a novel clamping mechanism.

Biologic tissues are complex materials that come in many forms and perform a variety of functions. They vary widely in composition and mechanical properties, and determination of the mechanical properties of tissues is of interest to those trying to engineer tissues to restore missing function. In performing experiments to characterize the mechanical properties of biologic tissues, there is no single solution to clamping tissues or tissue engineered constructs for mechanical testing. Various clamping techniques have been developed over the past few decades to address the difficulty of imposing appropriate boundary conditions on particular soft tissues during mechanical testing. Two criteria for a successful clamping mechanism are (i) prevention of test specimen slippage, and (ii) prevention of test specimen failure outside the gage region. Herein we present a novel clamping mechanism design developed for the mechanical testing of abdominal wall tissue as an example. This design incorporates pins with serrated clamps to successfully decrease the occurrence of test sample slippage while reducing imposed stress concentrations at the clamping sites. This design was evaluated by performing 40 uniaxial tensile tests on rat abdominal wall muscles using strain rates of 1% per second or 10% per second. Load and displacement data were acquired at the grips. The clamping area on the tissue sample was marked with India ink to track potential slippage of the sample during testing. Ultimate tensile strength and the corresponding stretch were calculated when the maximum load was achieved. With fine-tuning of the torque applied to the clamping grips, the success rate of the tensile tests reached over 90%.

Characterizing the non-linear mechanical behavior of native and biomimetic engineered tissues in 1D with physically meaningful parameters.

It is common practice to evaluate the mechanical performance of a scaffold for tissue engineering using concepts from linear elasticity theory (i.e. Young's modulus), or variations thereof, and uniaxial testing data. In some cases the non-linear nature of tissue stress-strain behavior has prompted development of empirical approaches to obtain a more comprehensive description of the observed mechanical behavior. Such approaches constitute improvements over singular stiffness measures but the lack of an appropriate non-linear theoretical foundation renders them somewhat arbitrary and potentially incomplete. Recently, a constitutive model for non-linear tissues was developed based on first principles in physics. The Freed-Rajagopal 1-D Fiber Model incorporates physically meaningful parameters that provide a unique and comprehensive characterization of non-linear tissue behavior for the class of tissues with strain limiting behavior in 1D. The physical interpretation that these parameters provide suggests they may serve as useful design targets for tissue engineering applications. In this study, the Freed-Rajagopal model is employed with conventional uniaxial mechanical testing data obtained from experiments with collagen scaffolds for hernia repair grafts and the healthy native tissue counterpart. Results from the Freed-Rajagopal analysis revealed that tissue-engineered constructs that qualify as "biomimetic" according to linear elasticity theory, or variations thereof, are not truly biomimetic, as they do not mimic the non-linear mechanical behaviors observed in their native tissue counterparts. Most importantly, the Freed-Rajagopal model was easy to employ (it can be done using a standard uniaxial testing system, with minimal additional effort) and revealed specific design improvements that could be targeted to improve the biofidelity of these constructs. A performance comparison with conventional non-linear models (including Fung's 1D Law and a one-dimensionalized version of the Holzapfel, Gasser, Ogden model), was then conducted and revealed the Freed-Rajagopal model produced results that correlated exceptionally well with experimental data and better describes material behavior at low strains as compared to competing models.

Ballistic thermal phonons traversing nanocrystalline domains in oriented polyethylene.

Thermally conductive polymer crystals are of both fundamental and practical interest for their high thermal conductivity that exceeds that of many metals. In particular, polyethylene fibers and oriented films with uniaxial thermal conductivity exceeding 50 [Formula: see text] have been reported recently, stimulating interest into the underlying microscopic thermal transport processes. While ab initio calculations have provided insight into microscopic phonon properties for perfect crystals, such properties of actual samples have remained experimentally inaccessible. Here, we report the direct observation of thermal phonons with mean free paths up to 200 nm in semicrystalline polyethylene films using transient grating spectroscopy. Many of the mean free paths substantially exceed the crystalline domain sizes measured using small-angle X-ray scattering, indicating that thermal phonons propagate ballistically within and across the nanocrystalline domains; those transmitting across domain boundaries contribute nearly one-third of the thermal conductivity. Our work provides a direct determination of thermal phonon propagation lengths in molecular solids, yielding insights into the microscopic origins of their high thermal conductivity.

Dispersal and reproductive careers of male mountain gorillas in Bwindi Impenetrable National Park, Uganda.

Dispersal is a key event in the life of an animal and it influences individual reproductive success. Male mountain gorillas exhibit both philopatry and dispersal, resulting in a mixed one-male and multimale social organization. However, little is known about the relationship between male dispersal or philopatry and reproductive careers in Bwindi mountain gorillas. Here we analyze data spanning from 1993 to 2017 on social groups in Bwindi Impenetrable National Park, Uganda to examine the proportion of males that disperse, age of dispersal, pathways to attaining alpha status, fate of dispersing males and philopatric males, and male tenure length as well as make comparisons of these variables to the Virunga mountain gorilla population. We report previously undocumented cases of dispersal by immature males and old males and we also observed the only known case of a fully mature male immigrating into a breeding group. We used genetic tracking of known individuals to estimate that a minimum of 25% of males that disperse to become solitary males eventually form new groups. No differences were found between the Bwindi and Virunga population in the age of male dispersal, the proportion of males that disperse, the age of alpha male acquisition, and dominance tenure length. The lack of differences may be due to small sample sizes or because the observed ecological variability does not lead to life history differences between the populations. Males in both populations follow variable strategies to attain alpha status leading to the variable one-male and multimale social organization, including dispersal to become solitary and eventually form a group, via group fissioning, usurping another alpha male, or inheriting the alpha position when a previous group leader dies.

Quadratic relationships between group size and foraging efficiency in a herbivorous primate.

The effect of feeding competition on foraging efficiency is an important link between ecological factors and the social organization of gregarious species. We examined the effects of group size on daily travel distances, activity budgets, and energy intake of mountain gorillas in Rwanda. We measured daily travel distances of five groups, activity budgets of 79 gorillas in nine groups, and energy intake data for 23 adult females in three groups over a 16-month period. Travel distances and the proportion of time spent traveling increased with size for most groups, which would be expected if their foraging efficiency is limited by intragroup feeding competition. However, travel distances and times decreased for the largest group, which also had higher energy intake rates than intermediate sized groups. The improved foraging efficiency of the largest group may be explained by advantages in intergroup contest competition. The largest group had much lower home range overlap than the other study groups which may be due to groups avoiding one another as a result of male mating competition. Collectively, our results indicate that intermediate sized groups had the lowest foraging efficiency and provide a new twist on the growing evidence of non-linear relationships between group size and foraging efficiency in primates.

Variation in the social organization of gorillas: Life history and socioecological perspectives.

A focus of socioecological research is to understand how ecological, social, and life history factors influence the variability of social organization within and between species. The genus Gorilla exhibits variability in social organization with western gorilla groups being almost exclusively one-male, yet approximately 40% of mountain gorilla groups are multimale. We review five ultimate causes for the variability in social organization within and among gorilla populations: human disturbance, ecological constraints on group size, risk of infanticide, life history patterns, and population density. We find the most evidence for the ecological constraints and life history hypotheses, but an over-riding explanation remains elusive. The variability may hinge on variation in female dispersal patterns, as females seek a group of optimal size and with a good protector male. Our review illustrates the challenges of understanding why the social organization of closely related species may deviate from predictions based on socioecological and life history theory.