Functional craniology and brain evolution: from paleontology to biomedicine

“Because organisms are not a collection of independent traits, but rather integrated entities (Moss and Young, 1960; Gould and Lewontin, 1979; Enlow and Hans, 1996), it is important to consider variation among spatially proximate features of the skull, and how longterm evolutionary trends may impact…”

“In this sense, brain metabolism and thermoregulation are good examples of biological functions that may be modeled based on simple mechanistic and geometric principles (Nelson and Nunneley, 1998; Van Leeuwen et al., 2000; Sukstanskii and Yablonskiy, 2006; Zhu et al., 2006; Bruner et al., 2011a,…”

“Brains are well known for being among the most energy-demanding organs of the body, burning large quantities of glucose for the development and maintenance of their structural and functional integrity (Mink et al., 1981; Aiello and Wheeler, 1995; Leonard et al., 2007).”

“(2007). Open Access Series of Imaging Studies (OASIS): cross-sectional MRI”

“In morphometrics it can be useful to separate variation (the actual distribution of a phenotype) and variability (the intrinsic possibility to vary, changing the distribution of the variation; Wagner and Altenberg, 1996).”

“On a smaller scale, local structures can be influenced by geometrical and physical properties at even the cellular level (Van Essen, 1997; Hilgetag and Barbas, 2005; Toro and Burnod, 2005).”

“REVIEW ARTICLE published: 02 April 2014 doi: 10.3389/fnana.2014.00019 Functional craniology and brain evolution: from paleontology to biomedicine”

“Interestingly, it also shows exceptional levels of energy management (Cavanna and Trimble, 2006; Margulies et al., 2009; Sotero and Iturria-Medina, 2011; Zhang and Li, 2012).”

“Relative to body size the human brain is very large, with a majority of this increase beginning during the Middle Pleistocene (Ruff et al., 1997; Rightmire, 2004).”

“…morphology, one of the main goals is to investigate the degree and patterns of integration within a given anatomical system, or alternatively, the separation of modules characterized by highly correlated traits (Cheverud, 1982, 1996; Mitteroecker and Bookstein, 2007; Wagner et al., 2007).”

“REVIEW ARTICLE published: 02 April 2014 doi: 10.3389/fnana.2014.00019 Functional craniology and brain evolution: from paleontology to biomedicine”

“Because organisms are not a collection of independent traits, but rather integrated entities (Moss and Young, 1960; Gould and Lewontin, 1979; Enlow and Hans, 1996), it is important to consider variation among spatially proximate features of the skull, and how longterm evolutionary trends may impact…”

“In the last two decades, growing attention has been placed on the relationship between evolutionary constraints and pathology, which has contributed to the development of perspectives in medicine that are based on an evolutionary foundation (Williams and Nesse, 1991).”

“In terms of cranial integration, the facial and neural blocks are partially separated, but both interact with the cranial base (Bastir et al., 2010).”

“An increase in absolute and relative brain size is arguably the main hallmark of the human evolutionary lineages, and is generally associated with evidence of increased cultural and behavioral complexity (see Sherwood et al., 2008).”

“As far as this can be observed in endocranial casts, the modern sulcal and gyral patterns at the frontal lobes can be observed in every human species, dating back 2 million years (Tobias, 1987; Holloway, 1995).”

“…of endocranial form and shape also provides evidence that modern humans are characterized by a species-specific upper bulging of the parietal areas (Bruner et al., 2003, 2011b; Bruner, 2004), and results are the same if we consider endocranial (bone) or cortical (brain) anatomical references.”

“In terms of cytoarchitecture, humans have specific cellular areas in the intra-parietal sulcus when compared with non-human primates (Orban et al., 2006).”

“Recent paleoneurological inferences, based on correlations between cranial and cerebral morphology, suggest that Neanderthals had relatively larger occipital lobes when compared with modern humans (Pearce et al., 2013).”

“This paper is funded by a Grant-in-Aid for Scientific Research on Innovative Areas “Replacement of Neanderthals by Modern Humans: Testing Evolutionary Models of Learning” from the Japanese Ministry of Education, Culture, Sports, Science, and Technology (#22101006).”

“Anthropology was one of the first fields applying such new toolkits (Slice, 2005), which were soon used to investigate integration of the human skull in ontogeny and evolution (e.g., Bookstein et al., 2003; Bastir and Rosas, 2005; Bastir et al., 2006).”

“In addition to being axially elongated, the eye of myopes is also ubiquitously larger (Zadnik et al., 1994; Ip et al., 2007; Lam et al., 2008), though it is still unknown why the eye exhibits these Frontiers in Neuroanatomy www.frontiersin.org April 2014 | Volume 8 | Article 19 | 6 characteristics…”

“…to body size (Schultz, 1940), the well-documented reduction in overall human size and robusticity that began 12,000 years ago (Carlson, 1976; Carlson and Van Gerven, 1977; Smith et al., 1985, 1986; Henneberg, 1988; Henneberg and Steyn, 1993; Lahr and Wright, 1996) would act to increase the…”

“…morphology, one of the main goals is to investigate the degree and patterns of integration within a given anatomical system, or alternatively, the separation of modules characterized by highly correlated traits (Cheverud, 1982, 1996; Mitteroecker and Bookstein, 2007; Wagner et al., 2007).”

“Because dimensions of the orbital margins and body mass are highly correlated (Kappelman, 1996), and a negative allometric relationship exists between the eye and orbit with respect to body size (Schultz, 1940), the well-documented reduction in overall human size and robusticity that began 12,000…”

“Preliminary data suggested that the volume of the human frontal lobes is in line with what we would expect for a primate of the same brain size (Semendeferi et al., 1997; Semendeferi and Damasio, 2000).”

“…evolutionary level, as brain temperature homeostasis may impose pervasive selective pressures and constraints on the evolution of species, and particularly on species-specific encephalization processes (Falk, 1990; Hofman, 2001, 2012; Caputa, 2004; Bruner et al., 2011a, 2012; Manger et al., 2013).”

“In terms of cranial integration, the facial and neural blocks are partially separated, but both interact with the cranial base (Bastir et al., 2010).”

“…as it has yet to be shown how convergence and eye strain associated with more reading and near vision work can permanently alter the shape of the eye, and no other environmental risk factors that alter ocular growth have been identified (Saw et al., 2006; Mutti and Zadnik, 2009; Mutti, 2011).”

“Preliminary data suggested that the volume of the human frontal lobes is in line with what we would expect for a primate of the same brain size (Semendeferi et al., 1997; Semendeferi and Damasio, 2000).”

“Since this time there has been a nearly twofold increase in cranial size, however, enlargement of the various structures that make up the brain housed within it have not increased isometrically (Rilling, 2006).”

“It is also found to occur earlier in life and at a higher frequency among Chinese schoolchildren compared to individuals of African or European descent (Lam et al., 1999; Ip et al., 2007, 2008).”

“Such spatial changes in parietal geometry are associated with an early postnatal stage in our species (Neubauer et al., 2009), which is absent in chimpanzees (Neubauer et al., 2010) and Neanderthals (Gunz et al., 2010).”

“On a smaller scale, local structures can be influenced by geometrical and physical properties at even the cellular level (Van Essen, 1997; Hilgetag and Barbas, 2005; Toro and Burnod, 2005).”

“…and functional changes at the cellular, histological and systemic levels, while severe and irreversible neural damage, coma, or even death of the individual may happen under hyperthermic conditions with brain temperatures above 40◦C (Kiyatkin, 2010; Bertolizio et al., 2011; Rango et al., 2012).”

“Anthropology was one of the first fields applying such new toolkits (Slice, 2005), which were soon used to investigate integration of the human skull in ontogeny and evolution (e.g., Bookstein et al., 2003; Bastir and Rosas, 2005; Bastir et al., 2006).”

“…its influence on craniofacial development throughout ontogeny, has been hypothesized to be associated with secondary problems such as choking, dental crowding, impacted dentition, as well as a reduction in olfactory and masticatory function (Ross, 1995; Lieberman et al., 2000; Ravosa et al., 2000).”

“Although the eyeball lies predominantly within the orbit, it does not directly influence orbital size in humans (Schultz, 1940; Chau et al., 2004), but rather eye growth keeps pace with growth of the brain (Salzmann, 1912; Todd et al.”

“More recently however, it has been shown that in the earliest stages of the disease, metabolic impairments, neuroanatomical, and histological changes, can be found in the medial parietal cortex (Jacobs et al., 2012, 2013; Doré et al., 2013; Huang et al., 2013).”

“, Jolles, J., and Verhey, F. R. J. (2013). Decreased grey matter diffusivity: a”

“In general, any geometrical correlations among brain and skull landmarks are necessary to understand paleoneurological changes, as well as for planning surgical operations (Ribas et al., 2006; Richtsmeier and Deleon, 2009).”

“Because of such fine biomechanical balance associated with morphogenesis Frontiers in Neuroanatomy www.frontiersin.org April 2014 | Volume 8 | Article 19 | 7 and histological properties, minor changes in the physical composition of the connective, osseous, or neural elements of the brain, can exert…”

“For example, volume of the human temporal lobes is larger than expected according to the proportions exhibited among living apes (Rilling and Seligman, 2002).”

“…outward from the orbit), has led some researchers to suggest that degree of refractive error be considered in studies of exophthalmia, as it was presumed that increased axial length of the eye in myopes causes it to protrude out from the orbit (Migliori and Gladstone, 1984; Quant and Woo, 1992).”

“…has been found to be the biggest contributor to the condition, and particularly among individuals over 12 years of age and of East-Asian ancestry (Ip et al., 2007), where myopia is so common that it affects 80–90% of individuals in some East-Asian populations (Goldschmidt et al., 2001; Lin et…”

“These spatial relationships between brain and braincase are of special interest for both paleoneurology and surgery. may induce structural and functional changes at the cellular, histological and systemic levels, while severe and irreversible neural damage, coma, or even death of the individual may…”

“The precuneus is a cortical element that is more sensitive to such constraints, and we must assume that in this sense brain shape changes associated with neurocranial globularity can affect this particular area to a greater extent.”

“In this sense, brain metabolism and thermoregulation are good examples of biological functions that may be modeled based on simple mechanistic and geometric principles (Nelson and Nunneley, 1998; Van Leeuwen et al., 2000; Sukstanskii and Yablonskiy, 2006; Zhu et al., 2006; Bruner et al., 2011a, 2012).”

“As a consequence, most comparisons do not reach the common statistical thresholds of significance (Barton and Venditti, 2013).”

“Instead, morphological changes in the parietal regions among hominoids are probably associated with deep and upper parietal volumes, like the precuneus or the intraparietal sulcus, which are largely involved in visuo-spatial integration (Bruner, 2010; Bruner et al., 2014).”

“…this sense, brain metabolism and thermoregulation are good examples of biological functions that may be modeled based on simple mechanistic and geometric principles (Nelson and Nunneley, 1998; Van Leeuwen et al., 2000; Sukstanskii and Yablonskiy, 2006; Zhu et al., 2006; Bruner et al., 2011a, 2012).”

“…that began 12,000 years ago (Carlson, 1976; Carlson and Van Gerven, 1977; Smith et al., 1985, 1986; Henneberg, 1988; Henneberg and Steyn, 1993; Lahr and Wright, 1996) would act to increase the percentage of the orbit occupied by the eye, and bring humans as a whole closer to the point at…”

“This is particularly relevant in cognitive sciences, considering current theories on the extended mind, in which the integration among brain, body, and environment are essential to generate our behavioral capacities (Iriki and Sakura, 2008; Malafouris, 2009; Iriki and Taoka, 2012).”

“As far as this can be observed in endocranial casts, the modern sulcal and gyral patterns at the frontal lobes can be observed in every human species, dating back 2 million years (Tobias, 1987; Holloway, 1995).”

“…forward projection of the greater sphenoid wings, and rotation of the posterior maxillary plane (McCarthy and Lieberman, 2001; Lieberman et al., 2002; Bastir et al., 2008; Cobb, 2008), as well as a reduction in anterior projection of the orbital margins and internal orbital depth (Masters, 2009).”

“A marked increase in their general proportions can be seen later in Neanderthals and modern humans, which show relatively wider prefrontal areas at the Broca’s cap when compared with other human species (Bruner and Holloway, 2010).”

“…this sense, brain metabolism and thermoregulation are good examples of biological functions that may be modeled based on simple mechanistic and geometric principles (Nelson and Nunneley, 1998; Van Leeuwen et al., 2000; Sukstanskii and Yablonskiy, 2006; Zhu et al., 2006; Bruner et al., 2011a, 2012).”

“…this sense, brain metabolism and thermoregulation are good examples of biological functions that may be modeled based on simple mechanistic and geometric principles (Nelson and Nunneley, 1998; Van Leeuwen et al., 2000; Sukstanskii and Yablonskiy, 2006; Zhu et al., 2006; Bruner et al., 2011a, 2012).”

“…movement of the anterior and middle cranial fossae, forward projection of the greater sphenoid wings, and rotation of the posterior maxillary plane (McCarthy and Lieberman, 2001; Lieberman et al., 2002; Bastir et al., 2008; Cobb, 2008), as well as a reduction in anterior projection of the orbital…”

“Beyond any possible coincidence, we should seriously evaluate the possibility that the vulnerability of the parietal areas, and the sensitivity to the processes of neurodegeneration associated with AD, could be a drawback of the complexity of our brain anatomy (Bruner and Jacobs, 2013).”

“REVIEW ARTICLE published: 02 April 2014 doi: 10.3389/fnana.2014.00019 Functional craniology and brain evolution: from paleontology to biomedicine”

“These changes in cerebral dimensions are integrated with modification to underlying basicranial and facial structures, and have been associated with an anteroposteriorly shorter face (Bookstein et al., 2003; Bastir and Rosas, 2009; Bastir et al., 2010).”

“In addition to being axially elongated, the eye of myopes is also ubiquitously larger (Zadnik et al., 1994; Ip et al., 2007; Lam et al., 2008), though it is still unknown why the eye exhibits these Frontiers in Neuroanatomy www.frontiersin.org April 2014 | Volume 8 | Article 19 | 6 characteristics…”

“As a consequence, the three endocranial fossae are not strictly integrated according to specific morphological schemes, but rather are influenced by distinct and local factors (Bruner and Ripani, 2008).”

“…human size and robusticity that began 12,000 years ago (Carlson, 1976; Carlson and Van Gerven, 1977; Smith et al., 1985, 1986; Henneberg, 1988; Henneberg and Steyn, 1993; Lahr and Wright, 1996) would act to increase the percentage of the orbit occupied by the eye, and bring humans as a whole…”

“Taking into account that such close contact is not observed in less encephalized hominids (Bookstein et al., 1999; Bruner and Manzi, 2005), we must assume that this form, involving greater interaction between these two anatomical components, has evolved in both lineages independently beginning…”

“…evolutionary level, as brain temperature homeostasis may impose pervasive selective pressures and constraints on the evolution of species, and particularly on species-specific encephalization processes (Falk, 1990; Hofman, 2001, 2012; Caputa, 2004; Bruner et al., 2011a, 2012; Manger et al., 2013).”

“A recent geometric analysis has shown that relative dimensions of the precuneus are largely responsible for these major patterns of variation (Bruner et al., 2014; Figure 4B).”

“Craniosynostoses and suture dynamics are other essential topics in both fields, being associated with morphogenetic factors involved in pathology and phylogeny (Di Ieva et al., 2013).”

“…evolutionary level, as brain temperature homeostasis may impose pervasive selective pressures and constraints on the evolution of species, and particularly on species-specific encephalization processes (Falk, 1990; Hofman, 2001, 2012; Caputa, 2004; Bruner et al., 2011a, 2012; Manger et al., 2013).”

“…as it has yet to be shown how convergence and eye strain associated with more reading and near vision work can permanently alter the shape of the eye, and no other environmental risk factors that alter ocular growth have been identified (Saw et al., 2006; Mutti and Zadnik, 2009; Mutti, 2011).”

“Though independent of any correlations with cognitive abilities or the primary selective processes behind such evolutionary changes (Alba, 2010; Manger et al., 2013), we know that big brains are expensive and difficult to handle in terms of ecology and anatomy.”

“J., and Lewontin, R. C. (1979). The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme.”

“Taking into account that such close contact is not observed in less encephalized hominids (Bookstein et al., 1999; Bruner and Manzi, 2005), we must assume that this form, involving greater interaction between these two anatomical components, has evolved in both lineages independently beginning…”

“…of the brain in a sample of adult modern humans, we can see that frontoparietal bulging is also the major source of intra-specific variability (Bruner et al., 2010), suggesting the probable role during human evolution of lines of least resistance, in which intra-specific variability…”

“Such frequency has been measured as “hypostotic scores,” and interpreted as a structural result of ontogenetic stress (Manzi et al., 2000).”

“…contributor to the condition, and particularly among individuals over 12 years of age and of East-Asian ancestry (Ip et al., 2007), where myopia is so common that it affects 80–90% of individuals in some East-Asian populations (Goldschmidt et al., 2001; Lin et al., 2004; Park and Congdon, 2004).”

“While heat production and active removal of heat by cerebral blood flow are correlated with overall cerebral dimensions (Karbowski, 2007, 2009; HerculanoHouzel, 2011), passive heat diffusion (i.e., conduction) within the brain mostly depends on cerebral shape (Bruner et al., 2011a, 2012).”

“Frontiers in Neuroanatomy www.frontiersin.org April 2014 | Volume 8 | Article 19 | 8 lack such features (Bruner and Pearson, 2013).”

“…it does not directly influence orbital size in humans (Schultz, 1940; Chau et al., 2004), but rather eye growth keeps pace with growth of the brain (Salzmann, 1912; Todd et al., 1940; Weale, 1982), and both are thought to be the product of pleiotropic gene control (Miller, 1992; Mak et al., 2006).”

“Although such differences are macroscopic, we still ignore the functional and evolutionary meaning of this specific change (Falk, 1993; Bruner et al., 2011a).”

“Interestingly, reduced visual acuity is also common among numerous dog breeds such as the Toy Poodle, Miniature Schnauzer, Pug, Rottweiler, Collie, and Labrador retriever (Murphy et al., 1992; Kubai et al., 2008; Williams et al., 2011).”

“Magnetic resonance is in fact recognized for its ability to visualize brain, but it can also be used to localize sutures, because of their connective composition (Cotton et al., 2005).”

“It is apparent that this same information is relevant in evolutionary biology as well as in biomedicine.”

“…1940), the well-documented reduction in overall human size and robusticity that began 12,000 years ago (Carlson, 1976; Carlson and Van Gerven, 1977; Smith et al., 1985, 1986; Henneberg, 1988; Henneberg and Steyn, 1993; Lahr and Wright, 1996) would act to increase the percentage of the orbit…”

“…and functional changes at the cellular, histological and systemic levels, while severe and irreversible neural damage, coma, or even death of the individual may happen under hyperthermic conditions with brain temperatures above 40◦C (Kiyatkin, 2010; Bertolizio et al., 2011; Rango et al., 2012).”

“During morphogenesis, imbalance among tissues may lead to non-pathological variants called “epigenetic traits” (Hauser and De Stefano, 1989; Manzi et al., 1996), and cranial morphogenesis is particularly sensitive to bone deposition and resorption associated with growth fields of osteoblasts and…”

“…contributor to the condition, and particularly among individuals over 12 years of age and of East-Asian ancestry (Ip et al., 2007), where myopia is so common that it affects 80–90% of individuals in some East-Asian populations (Goldschmidt et al., 2001; Lin et al., 2004; Park and Congdon, 2004).”

“More recently however, it has been shown that in the earliest stages of the disease, metabolic impairments, neuroanatomical, and histological changes, can be found in the medial parietal cortex (Jacobs et al., 2012, 2013; Doré et al., 2013; Huang et al., 2013).”

“However, a recent analysis of published data on eyeball volume, orbital volume, and refractive error in Chinese adults has indicated that relative size of the eye within the orbit may be an important predictor of myopia (Masters, 2012).”

“As previously mentioned, for the deep parietal areas we must consider that the precuneus, approaching the core of the brain in terms of localization, can be susceptible to high thermal loads because of its spatial position (Bruner et al., 2012).”

“…and functional changes at the cellular, histological and systemic levels, while severe and irreversible neural damage, coma, or even death of the individual may happen under hyperthermic conditions with brain temperatures above 40◦C (Kiyatkin, 2010; Bertolizio et al., 2011; Rango et al., 2012).”

“With this limitation in mind, there is no doubt that evolutionary anthropology, as natural history of the humankind, can represent an interesting and informative key to evaluate failures and successes of our species in a biomedical context.”

“This has been demonstrated most clearly in the Labrador retriever, where a significant genetic component exists (Black et al., 2008), and the greatest contributors to refractive error include a thinning lens and increased vitreous chamber depth (Mutti et al., 1999).”

“…it does not directly influence orbital size in humans (Schultz, 1940; Chau et al., 2004), but rather eye growth keeps pace with growth of the brain (Salzmann, 1912; Todd et al., 1940; Weale, 1982), and both are thought to be the product of pleiotropic gene control (Miller, 1992; Mak et al., 2006).”

“Interestingly, it also shows exceptional levels of energy management (Cavanna and Trimble, 2006; Margulies et al., 2009; Sotero and Iturria-Medina, 2011; Zhang and Li, 2012).”

“Interestingly, reduced visual acuity is also common among numerous dog breeds such as the Toy Poodle, Miniature Schnauzer, Pug, Rottweiler, Collie, and Labrador retriever (Murphy et al., 1992; Kubai et al., 2008; Williams et al., 2011).”

“Imbalances among these elements due to genetic or epigenetic factors are often associated with pathological and sub-pathological conditions, due to altered patterns of the ossification process (e.g., Morriss-Kay and Wilkie, 2005; Carter and Anslow, 2009; Richtsmeier and Flaherty, 2013).”

“…reconstruction of a modern human skull and endocast; b) orm, that is facial flattening and parietal bulging; c) endocranial outline, showing the g; d) the main pattern of intra-specific adult brain form variation in modern humans is Bruner, 2004; Bruner et al., 2004; Bruner et al., 2014a,b).”

“…of the Parietal Lobes 141Brain Behav Evol 2018;91:136–147 DOI: 10.1159/000488889 cranial heat exchange, and the bulging of the parietal region characteristic of modern humans can hence contribute to increase the heat load of the corresponding deeper cerebral areas [Bruner et al., 2012, 2014a].”

“These are also the only human species with the frontal lobes located entirely over the orbits (Bruner et al., 2014), but the functional implications of these findings are unclear (Balzeau et al., 2014; Bruner, 2017).”

“…the precuneus is also particularly active in metabolic terms (Cavanna and Trimble 2006; Sotero and IturriaMedina 2011), suggesting that its evolutionary changes may have been associated with thermal constraints due to the brain’s energy consumption and heat release (Bruner et al. 2014a).”

“In fact, Neanderthals and modern humans, although sharing a similar brain size, had different heat distribution patterns because of their different brain form (Bruner et al., 2014).”

“In a previous analysis using the same sample, we showed that the main pattern of morphological variation for the midsagittal brain section was associated with relative dilation/contraction of the precuneus (Bruner et al., 2014a).”

“Previousstudiesofancient crania often interpreted the shapedifferences as evidenceof relative size alterations of different cortical functional domains [5, 27].”

“Orbits are anterior to the braincase in chimps, inferior to the frontal lobes in Neandertals and modern humans, and in an intermediate position in archaic humans (Bruner et al., 2014a; Beaudet and Bruner, 2017; Figure 1).”

“All of these changes (contiguity between orbits and frontal lobes, reduced facial block, and enlarged temporal lobes) involve a spatial conflict between endocranial and orbital areas during growth and development, potentially leading to a deformation of the eye bulb and vision defects among more recent modern humans (…”

“Because of the many structural and functional relationships in the human parieto-occipital block, morphogenetic constraints may be expected and can even explain epigenetic hypostotic features associated with the Neanderthal’s braincase development (Bruner 2014).”

“This result suggests that functional matrices nearby, such as the brain, do not constrain the establishment of their articulations, even though the shape and orientation of the cranial base in primates is highly integrated to facial size and cranial encephalization (Bastir et al. 2010; Bruner et al. 2014).”

“The vascular system transports nutrients, oxygen, and hormones, and it is also involved in thermoregulation and other functions related to the maintenance of homeostasis (Moss and Young, 1960; Enlow, 1968; Bruner and Sherkat, 2008; Bruner et al., 2009, 2014; Percival and Richtsmeier, 2013).”

“Furthermore, preliminary comparisons between cranial and brain landmarks suggests that there is an important level of independence between cranial and brain boundaries, and the former are hence not necessarily a good proxy for estimating brain proportions (Bruner et al., 2014).”

“…projection of the temporal lobes behind the orbits (Bastir et al. 2008), would be expected to cause ocular distortion, and specifically axial elongation of the eye, as it shifts more toward the narrowing concave rim of the anterior orbital margins in recent human evolution (Bruner et al. 2014).”

“Digital data promote new methods in quantitative analysis of endocranial shape, notably through geometric morphometric methods (e.g., Bruner, 2004; Gunz, 2015; Bruner et al., 2003; Neubauer et al., 2010) and registration of surfaces from the correspondence of anatomical landmarks (e.”

“The middle meningeal artery is particularly developed on the parietal surface, which is likely a crucial region of heat interchange (Bruner & Sherkat, 2008; Bruner et al. 2014).”

“To achieve detailed inferences of brain morphology based on cranial shape, more microscopic correspondences between cranial and brain morphology must also be explored and investigated in future studies (Bruner et al., 2014).”

“Variations in the local and global proximity may influence physiological features that are sensitive to spatial distribution, like speed of communication, thermal exchange or biomechanical strains (Bruner et al., 2014).”

“There is such a fine biomechanical balance associated with morphogenesis that small changes in the biochemistry or structure of connective, osseous or neural elements of the brain can exert direct influence on the spatial relationship of endocranial components (Bruner et al. 2014).”

“However, human evolutionary lineages are marked by a unique increase in absolute and relative brain size commonly associated with increased behavioral, sociocultural, and linguistic complexity (e.g., Bruner 2004; Bruner et al. 2014; Sherwood et al. 2008).”

“It has a notable metabolic load and thermal load, and is associated with metabolic impairments and vulnerability to neurodegenerative processes like Alzheimer disease [Zhang and Li, 2012; Bruner and Jacobs, 2013; Bruner et al., 2014a].”

“…in the case of the cranial vault, according to the cranial functional matrix hypothesis, the brain is more likely to mold the skull due to this region's lower morphogenetic complexity (Bruner, 2015; Bruner et al., 2014; Moss & Young, 1960; Richtsmeier & Flaherty, 2013; Schuurman & Bruner, 2023).”

“The brain's spatial arrangement is also impacted by structural and functional interactions with the surrounding braincase, specifically from the physical interaction between soft and hard tissues (Bruner et al., 2014; Moss & Young, 1960).”

“It has been also hypothesized that the complexity and topology of these vessels might be related to the balance between heat production and dissipation, which is influenced by changes in both brain size and shape (Bruner et al., 2011, 2012, 2014).”

“There is no doubt that frontal areas changed their spatial relationships during human evolution: in Neanderthals and modern humans, they are positioned onto the orbital roof, introducing some constraints associated with the relationships between the brain and the upper facial block (Bruner et al. 2014).”

“Facial skeleton retraction is a major characteristic of B individuals, associated with a decrease in skull length (anterior to posterior) (Bruner et al., 2014), causing variations in the cp spatial position, such as the g.”

“The morphology of the human brain and skull reflects, besides function, intrinsic and extrinsic spatial constraints (Bruner et al., 2014; Hofman, 2012; Moss & Young, 1960).”

“The Homo endocasts support these hypotheses to study the traces left by the brain inside the skull (Bruner, 2017; Bruner et al., 2014; Neubauer, 2014; Pestana et al., 2023).”

“The vein of Galen (1) is expected inferior to the splenium (2), draining to the straight sinus (3).”

“These cortical areas have been hypothesized to undergo an expansion in Homo sapiens (Bruner, Preuss, et al. 2017; Pereira-Pedro et al. 2020), with a possible increase in the thermal load of the underlying anatomical regions (Bruner et al. 2014).”

“Candidates include encephalization with brain reorganization, facial size reduction, and posture and locomotion, which would have influenced the basicranial morphology, metabolism, and cranial hemodynamics (Bastir et al., 2010, 2011; Bruner et al., 2014; Neaux et al., 2018; Seymour et al., 2019).”

“Such changes in brain proportions probably involved deep parietal elements, such as the intraparietal sulcus and the precuneus (Bruner et al., 2014).”

“13 In this paper, there was found a low positive correlation between ETD and cranium measurements which are NA-OP ( r = 0.391), NA-BA ( r = 0.386), NA-dorsum sella ( r = 0.326), and PR-BA ( r = 0.366).”

“At the time of birth, the sphenoid sinuses are present but not yet aerated, with a total volume not exceeding 2 mm 3 —comparable in size to a small pea—representing cavities continuous with the spheno-ethmoidal recesses [25,40,50,51,53].”