Moving beyond the adaptationist paradigm for human evolution, and why it matters.

The Journal of Human Evolution (JHE) was founded 50 years ago when much of the foundation for how we think about human evolution was in place or being put in place, providing the main framework for how we consider our origins today. Here, we will explore historical developments, including early JHE outputs, as they relate to our understanding of the relationship between phenotypic variation and evolutionary process, and use that as a springboard for considering our current understanding of these links as applied to human evolution. We will focus specifically on how the study of variation itself has shifted us away from taxonomic and adaptationist perspectives toward a richer understanding of the processes shaping human evolutionary history, using literature searches and specific test cases to highlight this. We argue that natural selection, gene exchange, genetic drift, and mutation should not be considered individually when considering the production of hominin diversity. In this context, we offer suggestions for future research directions and reflect on this more complex understanding of human evolution and its broader relevance to society. Finally, we end by considering authorship demographics and practices in the last 50 years within JHE and how a shift in these demographics has the potential to reshape the science of human evolution going forward.

Morphological integration in the hominid midfoot.

The calculation of morphological integration across living apes and humans may provide important insights into the potential influence of integration on evolutionary trajectories in the hominid lineage. Here, we quantify magnitudes of morphological integration among and within elements of the midfoot in great apes and humans to examine the link between locomotor differences and trait covariance. We test the hypothesis that the medial elements of the great ape foot are less morphologically integrated with one another compared to humans based on their abducted halluces, and aim to determine how adaptations for midfoot mobility/stiffness and locomotor specialization influence magnitudes of morphological integration. The study sample is composed of all cuneiforms, the navicular, the cuboid, and metatarsals 1-5 of Homo sapiens (n = 80), Pan troglodytes (n = 63), Gorilla gorilla (n = 39), and Pongo sp. (n = 41). Morphological integration was quantified using the integration coefficient of variation of interlandmark distances organized into sets of a priori-defined modules. Magnitudes of integration across these modules were then compared against sets of random traits from the whole midfoot. Results show that all nonhuman apes have less integrated medial elements, whereas humans have highly integrated medial elements, suggesting a link between hallucal abduction and reduced levels of morphological integration. However, we find considerable variation in magnitudes of morphological integration across metatarsals 2-5, the intermediate and lateral cuneiform, the cuboid, and navicular, emphasizing the influence of functional and nonfunctional factors in magnitudes of integration. Lastly, we find that humans and orangutans show the lowest overall magnitudes of integration in the midfoot, which may be related to their highly specialized functions, and suggest a link between strong diversifying selection and reduced magnitudes of morphological integration.

Skull variation in Afro-Eurasian monkeys results from both adaptive and non-adaptive evolutionary processes.

Afro-Eurasian monkeys originated in the Miocene and are the most species-rich modern primate family. Molecular and fossil data have provided considerable insight into their evolutionary divergence, but we know considerably less about the evolutionary processes that underlie these differences. Here, we apply tests developed from quantitative genetics theory to a large (n > 3000) cranio-mandibular morphometric dataset, investigating the relative importance of adaptation (natural selection) and neutral processes (genetic drift) in shaping diversity at different taxonomic levels, an approach applied previously to monkeys of the Americas, apes, hominins, and other vertebrate taxa. Results indicate that natural selection, particularly for differences in size, plays a significant role in diversifying Afro-Eurasian monkeys as a whole. However, drift appears to better explain skull divergence within the subfamily Colobinae, and in particular the African colobine clade, likely due to habitat fragmentation. Small and declining population sizes make it likely that drift will continue in this taxon, with potentially dire implications for genetic diversity and future resilience in the face of environmental change. For the other taxa, many of whom also have decreasing populations and are threatened, understanding adaptive pressures similarly helps identify relative vulnerability and may assist with prioritising scarce conservation resources.

The activity patterns of nonworking and working sled dogs.

There are limited studies investigating the combined effects of biological, environmental, and human factors on the activity of the domestic dog. Sled dogs offer a unique opportunity to examine these factors due to their close relationship with handlers and exposure to the outdoors. Here, we used accelerometers to measure the activity of 52 sled dogs over 30 days from two locations in Canada. The two locations differ in the working demands of dogs, therefore we used linear mixed effects models to assess how different factors impact daytime and nighttime activity of working versus nonworking dogs. During the daytime, we found that males were more active than females among nonworking dogs and younger dogs were more active than older dogs among working dogs. Alaskan huskies had higher activity levels than non-Alaskan husky breeds in working sled dogs during the day. Nonworking dogs were slightly more active during colder weather, but temperature had no effect on working dogs' activity. The strongest predictor of daytime activity in working dogs was work schedule. These results indicate that the influence of biological factors on activity varied depending on dogs' physical demands and human activity was the most powerful driver of activity in working dogs.

Ovariectomized rat model and shape variation in the bony labyrinth.

Postmenopausal osteoporosis is a serious concern in aging individuals, but has not been explored for its potential to alter the shape of the inner ear by way of increased remodeling in the otic capsule. The otic capsule, or bony labyrinth, is thought to experience uniquely limited remodeling after development due to high levels of osteoprotegerin. On this basis, despite the widespread remodeling that accompanies osteoporosis, we hypothesize that both the shape and volume of the semicircular canals will resist such changes. To test this hypothesis, we conducted three-dimensional geometric morphometric shape analysis on microcomputed tomographic data collected on the semicircular canals of an ovariectomized (OVX) rat model. A Procrustes ANOVA found no statistically significant differences in shape between surgery and sham groups, and morphological disparity testing likewise found no differences in shape variation. Univariate testing found no differences in semicircular volume between OVX and control groups. The range of variation in the OVX group, however, is greater than in the sham group but this difference does not reach statistical significance, perhaps because of a combination of small effect size and low sample size. This finding suggests that labyrinthine shape remains a tool for assessing phylogeny and function in the fossil record, but that it is possible that osteoporosis may be contributing to intraspecific shape variation in the bony labyrinth. This effect warrants further exploration at a microstructural level with continued focus on variables related to remodeling.

Effects of foot progression angle on knee mechanics during an anticipated cutting task: A statistical parametric mapping approach.

Cutting is considered a "high-risk" movement for anterior cruciate ligament (ACL) injuries. It has been established that sex differences exist during cutting, placing females at greater ACL injury risk. Foot progression angle (FPA) during landing has been shown to influence lower extremity mechanics, yet little is known how FPA influences mechanics during cutting. The purpose of this study was to compare two FPA conditions during cutting between males and females. Twenty-four males and females were tested using two FPA conditions: toe-in 15° (TI15) and toe-out 15° (TO15). Right knee joint kinematic and kinetic variables were measured using a motion capture system and force plate. Five successful trials were collected and compared between FPA conditions. One-dimensional statistical parametric mapping was used to assess changes in knee mechanics between males and females over the entire stance phase. The only sex × FPA effect found was knee flexion angle. Females cutting at TI15 had decreased knee flexion angle compared TO15 (p = 0.019). Significant sex main effects included knee abduction and rotation angles, and knee flexion and rotation moments. Significant FPA main effects included knee flexion, abduction and rotation angles. The results show cutting with a toe-in FPA of 15° is enough to induce changes in knee abduction angle while cutting with 15° toe-out FPA influenced knee flexion and rotation angles. These data suggest that different cutting FPAs may be influential on known ACL injury risk variables. However, more research is warranted on cutting FPA before FPA is targeted as part of ACL injury prevention protocols.

The hybrid skull of the eastern coyote (Canis latrans var.): Nonmetric traits and craniomandibular shape.

The increasing awareness that hybridization, and resultant gene flow, plays a major role in animal diversification has led to a growing number of studies that have focused on assessing the morphological consequences of this process. Analyses of mammalian hybrids have identified skeletal effects of hybridization, including a suite of anomalous dental and sutural traits on the skull that are present at high frequencies in hybrid populations. These studies have also detected consistent patterns of morphological shape and size differences between hybrids and parental taxa across a wide variety of organisms. However, more research is required to understand the universality of these traits and shape/size differences. Building on these previous studies, a sample of genetically determined canid hybrids was examined, specifically the eastern coyote (Canis latrans var.), a hybrid between coyotes, wolves, and dogs, to test whether this group exhibits a comparable pattern of anomalous nonmetric characters, and to assess differences in craniomandibular shape and size. First, specimens of C. latrans var., C. latrans, and C. lupus were scored for anomalous traits, including supernumerary and rotated teeth, dental crowding, and sutural anomalies. Geometric morphometric analyses were then conducted on a subset of these individuals to explore craniomandibular size and shape variation, as well as allometry. The results are largely consistent with other studies, indicating that the incidence of dental anomalies, dental crowding, and sutural anomalies is significantly higher in hybrids. However, differences are not significant for supernumerary teeth. The exploration of morphometric variation identifies intermediate morphology in the hybrids, and some indication of greater morphological variability in the mandible. When these results are combined with previous studies, they suggest that skeletal signatures of hybridization are common to different mammalian taxa across multiple generations; however, some traits such as supernumerary teeth may be lost after a few generations.

Common evolutionary patterns in the human nasal region across a worldwide sample.

OBJECTIVES: Variation in the external nasal region among human populations has long been proposed in the literature to reflect adaptations to facilitate thermoregulation, air conditioning, and moisture retention in local climates and environments. More specifically, adaptations in populations living in colder climates have often been assumed due to correlational relationships found between variation in the nasal region and climatic variables. Here, we test this hypothesis by applying a quantitative genetics approach based on the Lande model to assess whether variation in the nasal region can be explained by random neutral processes (e.g., genetic drift) or if non-random forces (i.e., adaptation) have contributed significantly to its diversity. MATERIALS AND METHODS: A mixed-sex sample representing 28 population groups from Howells' craniometric dataset were analyzed (n = 2504). Twenty standard measurements were chosen to reflect the external skeletal morphology of the nasal region. We apply statistical tests developed from evolutionary quantitative genetics theory to analyze patterns of within- and between-population divergence under a null hypothesis of genetic drift. RESULTS: This study finds a rejection of genetic drift in all analyses, across tests that involve all 28 populations, exclusively cold-climate populations, and with cold-climate populations excluded, indicating that non-random evolutionary forces have contributed significantly to variation in the nasal region overall. DISCUSSION: These results show that nasal region adaptation is not exclusive to cold-climate populations, which have often been implicated in the literature to drive nasal variation, instead suggesting that the propensity for adaptation in the nasal region is shared among all human populations.

Overground Walking Biomechanics of Dissatisfied Persons With Total Knee Replacements.

Patient dissatisfaction following total knee replacement (TKR) procedures is likely influenced by both subjective and objective aspects. Increased pain and reduced performance on clinical tests have been shown in persons who are dissatisfied with the outcome of their surgery. However, it is unknown how overground walking kinematics and kinetics might differ in the dissatisfied versus satisfied patients following TKR surgery. This study compared the lower-extremity walking kinematics and kinetics of patients dissatisfied with their TKR to that of satisfied patients and healthy controls. Thirty nine subjects completed walking trials, including nine dissatisfied and 15 satisfied TKR patients and 15 healthy controls. A 2 × 3 repeated -measures analysis of variance was used to assess differences between groups and limbs (P < .05). Dissatisfied persons showed significantly reduced loading-response and push-off peak vertical ground reaction forces, flexion range of motion, loading-response extension moments, and loading-response abduction moments compared to the controls. Peak loading-response and push-off vertical ground reaction forces and flexion range of motion were reduced in the replaced limb of dissatisfied patients compared with their nonreplaced limb. Push-off plantar flexion moments were reduced in the dissatisfied patients compared with the satisfied and healthy controls. Dissatisfied patients also reported increased knee joint pain and reduced preferred gait speed. Moreover, dissatisfied patients experienced mechanical limb asymmetries not present in those satisfied with their surgery result. Thus, patients dissatisfied with their total knee replacement outcome were found to be experiencing significant negative physiological changes.

Increased Ankle Range of Motion Reduces Knee Loads During Landing in Healthy Adults.

Decreased dorsiflexion range of motion (DROM) can be modified using static stretching and joint mobilizations and may attenuate known knee anterior cruciate ligament injury risk factors. It is not known how these interventions compare to each other and how they improve knee landing mechanics. This study's purpose was to determine the immediate effects of static stretching and joint mobilization interventions on DROM measurement changes and right-leg drop jump knee landing mechanics. Eighteen females and 7 males, all recreationally active, completed 2 study sessions. Active and passive DROM, the weight-bearing lunge test, the anterior reach portion of the Star Excursion Balance Test, and a right-leg drop jump landing task were completed before and after the intervention. Change in DROM (ΔDROM) was calculated for DROM assessments between preintervention and postintervention. Pairwise dependent t tests determined no differences in ΔDROM between interventions, and statistical parametric mapping determined increased knee flexion (P = .004) and decreased anterior shear force (P = .015) during landing after both interventions. Increased DROM improves sagittal plane displacement and loading at the knee. Stretching may be a more feasible option in a healthy population for those wanting to maintain range of motion and decrease knee injury risk without physical therapist involvement.

Lower extremity muscle contributions to ACL loading during a stop-jump task.

Landing is considered a high-risk movement, especially landings from a stop-jump task, as they are often associated with lower extremity injuries, such as anterior cruciate ligament injuries (ACL). Females demonstrate lower extremity landing mechanics that often place them at a larger risk of injury compared to their male counterparts. While efforts have been made to understand lower extremity mechanics during stop-jump landings, little is known regarding the musculature function during these tasks and how they may influence ACL loading. Understanding lower extremity muscle contributions to ACL loading (FACL) may give insight to improving injury prevention protocols. Ten healthy, recreationally active females completed five trials of an unanticipated stop-jump task. Right leg kinematics, kinetics, and electromyography data were collected with three-dimensional motion capture, force plates, and electromyography sensors, respectively. Modified musculoskeletal models were scaled based on participant-specific anthropometrics, and muscle forces were obtained using static optimization. An induced acceleration analysis combined with a previously established mathematical ACL loading model was used to calculate lower extremity muscle contribution to FACL. The vastus lateralis, vastus intermedius, vastus medials, biceps femoris long head, semimembranosus, and soleus were found to be the primary contributors to FACL, with the vastus lateralis being the largest contributor. These data suggest that muscles traditionally known as ACL unloaders may in certain conditions load the ACL. These results also suggest that future injury prevention protocols should target muscles specifically to mitigate the influence the vastus lateralis has on ACL loading.

Early life malnutrition and fluctuating asymmetry in the rat bony labyrinth.

Maternal malnutrition during gestation and lactation is known to have adverse effects on offspring. We evaluate the impact of maternal diet on offspring bony labyrinth morphology. The bony labyrinth develops early and is thought to be stable to protect vital sensory organs within. For these reasons, bony labyrinth morphology has been used extensively to assess locomotion, hearing function, and phylogeny in primates and numerous other taxa. While variation related to these parameters has been documented, there is still a component of intraspecific variation that is unexplained. Although the labyrinthine developmental window is small, it may provide the opportunity for developmental instability to produce corresponding shape differences, as measured by fluctuating asymmetry (FA). We hypothesized that (a) offspring with poor maternal diet would exhibit increased FA, but (b) no unilateral shape difference. To test these hypotheses, we used two groups of rats (Rattus norvegicus; Crl:WI[Han] strain), one control group and one group exposed to a isocaloric, protein-restricted maternal diet during gestation and suckling. Individuals were sampled at weaning, sexual maturity, and old age. A Procrustes analysis of variance identified statistically significant FA in all diet-age subgroups. No differences in level of FA were identified among the subgroups, rejecting our first hypothesis. A principal components analysis identified no unilateral shape differences, supporting our second hypothesis. These results indicate that bony labyrinth morphology is remarkably stable and likely protected from a poor maternal diet during development. In light of this result, other factors must be explored to explain intraspecific variation in labyrinthine shape.

Type of unanticipated stimulus affects lower extremity kinematics and kinetics during sidestepping.

Including an unanticipated stimulus has significant impacts on lower extremity biomechanics during dynamic movements. It is unknown how a live, human defender affects lower extremity biomechanics. The purpose of this study was thus to determine the effects of two types of unanticipated stimuli (visual stimulus; defensive opponent) on lower extremity kinematics and kinetics in males and females during 45° sidestepping trials. Eight males and eight females completed two unanticipated stimuli sidestepping conditions. Numeric visual analog scales for reaction difficulty and movement realism were collected and analysed using a 2 × 2 mixed-model ANOVA. Three-dimensional hip, knee, and ankle kinematics and kinetics were measured during the stance phase of the sidestep and analysed using statistical parametric mapping. Participants reported greater difficulty and less realistic movements with the visual stimulus. Unanticipated stimulus main effects were observed for knee abduction angle, and hip extension and adduction, and knee extension and adduction moments. Sex main effects were observed for hip flexion, hip abduction, and ankle dorsiflexion angles, as well as hip abduction, ankle plantarflexion and ankle eversion moments and vertical ground reaction forces. Participants responded differently to two unanticipated stimuli. Careful consideration should be used when determining the type of unanticipated stimulus used.

Hybridization in human evolution: Insights from other organisms.

During the late Pleistocene, isolated lineages of hominins exchanged genes thus influencing genomic variation in humans in both the past and present. However, the dynamics of this genetic exchange and associated phenotypic consequences through time remain poorly understood. Gene exchange across divergent lineages can result in myriad outcomes arising from these dynamics and the environmental conditions under which it occurs. Here we draw from our collective research across various organisms, illustrating some of the ways in which gene exchange can structure genomic/phenotypic diversity within/among species. We present a range of examples relevant to questions about the evolution of hominins. These examples are not meant to be exhaustive, but rather illustrative of the diverse evolutionary causes/consequences of hybridization, highlighting potential drivers of human evolution in the context of hybridization including: influences on adaptive evolution, climate change, developmental systems, sex-differences in behavior, Haldane's rule and the large X-effect, and transgressive phenotypic variation.

Increased knee loading in stair ambulation in patients dissatisfied with their total knee replacement.

BACKGROUND: Total knee replacement patients have shown reductions in knee flexion range of motion, knee extensor moments, and gait speed during stair ascent and stair descent. However, it is unknown how patients dissatisfied with their total knee replacement differ from those who are satisfied during more difficult activities such as stair negotiation. Therefore, the purpose of this study was to compare knee biomechanics of patients who are dissatisfied with their joint replacement to those who are satisfied and healthy participants during stair negotiation. METHODS: Nine dissatisfied, fifteen satisfied patients and fifteen healthy participants participated, completing stair ascent and descent trials on an instrumented staircase. A 2 × 3 ANOVA was used to analyze biomechanical differences between groups and limbs during both activities. FINDINGS: The dissatisfied group showed reduced 2nd peak vertical GRF (P ≤ 0.0040) and loading-response knee extension moments (P ≤ 0.0041) in their operated limb compared to their non-operated limb and to satisfied and healthy groups during stair ascent. First peak vertical GRF (P < 0.0088) and both loading-response (P < 0.0117) and push-off abduction moments (P < 0.0028) showed reduced values in operated limbs compared to non-operated limbs for all groups. During stair descent, the dissatisfied group showed reduced loading-response and push-off knee extension moments (P ≤ 0.006) in their operated limb compared to their non-operated limb and the healthy group. The loading-response knee extension (P < 0.0379) and abduction moments (P ≤ 0.0048) were also reduced in the dissatisfied group compared to the satisfied group. INTERPRETATION: Patients who were dissatisfied showed asymmetrical loading of the knees in conjunction, which may have contributed to their dissatisfaction.

Hinged ankle braces do not alter knee mechanics during sidestep cutting.

Lateral ankle sprains are common injuries in quick, dynamic movements and are caused by rapid ankle inversion. Ankle braces are used to reduce ankle inversion, while allowing normal plantar and dorsiflexion ranges of motion. Knee injuries, such as anterior cruciate ligament injuries, are also common in dynamic movements. It is important to understand how ankle braces affect injury risk at other proximal joints. There is limited and conflicting results on how ankle braces affect knee mechanics during these types of movements. Additionally, it is unknown if sex differences exist when using an ankle brace. Therefore, the purpose of this study was to determine the effects of a hinged ankle brace and sex during a 45° cutting movement. Three-dimensional kinematics and ground reaction forces were collected using a motion capture system and force plate on ten men and eight women during cutting trials. 2 × 2 repeated measures ANOVAs were used to detect differences in ground reaction forces, as well as knee and ankle kinematics between brace conditions and sex (p < 0.05). The brace condition exhibited greater initial contact ankle dorsiflexion (p = 0.011), decreased peak ankle inversion (p < 0.01), and increased vertical loading rate (p = 0.040). Females performed the cutting movement with less initial contact (p = 0.019) and peak knee flexion (p = 0.023) compared to males. Ankle bracing had no impact on the observed sex differences. Females exhibited decreased knee flexion compared to males, which has been well documented in the literature. The use of an ankle braces reduced ankle injury risk variables while not adversely impacting knee mechanics during a 45° sidecutting movement.

Comparing Anterior Cruciate Ligament Injury Risk Variables Between Unanticipated Cutting and Decelerating Tasks.

To examine the relationship between anterior cruciate ligament injury risk factors in unanticipated cutting and decelerating. Three-dimensional kinematics and ground reaction forces were collected on 11 females (22 [2] y, 1.67 [0.08] m, and 68.5 [9.8] kg) during 2 unanticipated tasks. Paired samples t tests were performed to compare dependent variables between tasks. Spearman rank correlation coefficients were calculated to analyze the relationship between peak internal knee adduction moment and peak anterior tibial shear force (ASF) during 2 unanticipated tasks. Significantly greater knee abduction angles, peak knee adduction moments, and peak ASFs were observed during cutting (P ≤ .05). A strong positive correlation existed between decelerating ASF and cutting ASF (ρ = .67), while correlations between decelerating knee adduction moment and cutting knee adduction moment and decelerating ASF and cutting knee adduction moment were not significant. In situations where time management is a necessity and only one task can be evaluated, it may be more appropriate to utilize an unanticipated cutting task rather than an unanticipated deceleration task because of the increased knee adduction moment and ASF. These data can help future clinicians in better designing more effective anterior cruciate ligament injury risk screening methods.

Morphological integration of anatomical, developmental, and functional postcranial modules in the crab-eating macaque (Macaca fascicularis).

OBJECTIVES: Integration and modularity reflect the coordinated action of past evolutionary processes and, in turn, constrain or facilitate phenotypic evolvability. Here, we analyze magnitudes of integration in the macaque postcranium to test whether 20 a priori defined modules are (1) more tightly integrated than random sets of postcranial traits, and (2) are differentiated based on mode of definition, with developmental modules expected to be more integrated than functional or anatomical modules. MATERIALS AND METHODS: The 3D morphometric data collected for eight limb and girdle bones for 60 macaques were collated into anatomical, developmental, and functional modules. A resampling technique was used to create random samples of integration values for each module for statistical comparison. RESULTS: Our results found that not all a priori defined modules were more strongly integrated than random samples of postcranial traits and that specific types of modules did not present consistent patterns of integration. Rather, girdle and joint modules were consistently less integrated than limb modules, and forelimb elements were less integrated than hindlimbs. DISCUSSION: The results suggest that morphometrically complex modules tend to be less integrated than simple limb bones, irrespective of the number of available traits. However, differences in integration of the fore- and hindlimb more likely reflects the multitude of locomotory, feeding, and social functions involved. It remains to be tested whether patterns of integration identified here are primate universals, and to what extent they vary depending on phylogenetic or functional factors.

Evolutionary processes shaping diversity across the Homo lineage.

Recent fossil finds have highlighted extensive morphological diversity within our genus, Homo, and the co-existence of a number of species. However, little is known about the evolutionary processes responsible for producing this diversity. Understanding the action of these processes can provide insight into how and why our lineage evolved and diversified. Here, we examine cranial and mandibular variation and diversification from the earliest emergence of our genus at 2.8 Ma until the Late Pleistocene (0.126-0.0117 Ma), using statistical tests developed from quantitative genetics theory to evaluate whether stochastic (genetic drift) versus non-stochastic (selection) processes were responsible for the observed variation. Results show that random processes can account for species diversification for most traits, including neurocranial diversification, and across all time periods. Where selection was found to shape diversification, we show that: 1) adaptation was important in the earliest migration of Homo out of Africa; 2) selection played a role in shaping mandibular and maxillary diversity among Homo groups, possibly due to dietary differences; and 3) Homo rudolfensis is adaptively different from other early Homo taxa, including the earliest known Homo specimen. These results show that genetic drift, and, likely, small population sizes were important factors shaping the evolution of Homo and many of its novel traits, but that selection played an essential role in driving adaptation to new contexts.