At Holloway’s

Ralph Holloway is at the Columbia University (New York) since 1964. More than half century dedicated to paleoneurology, brain evolution, fossils and endocasts. Some weeks ago I was visiting his laboratory, an amazing place, full of books, experience, and history. And collections. Endocasts are everywhere, witnessing at once the evolution of the human brain and the evolution of the moulding techniques. So I took the opportunity to ask Ralph some quick comments for this blog …

Why do we still need physical casts? (but do we?)

I think for the most part, we can do very well with virtual endocasts (as long as these don’t get hacked…), although these can never provide the same haptic experience as a true cast, even if it is a good 3D print. At the moment, I am working on LES1, Homo naledi, and while I have a 3D print from scan data of the endocast surface, and good images provided by Heather Garvin, I am making an endocast from a 3D print of the cranial portion, as I need the best resolution I can get of the occipital portion in particular. Even micro-CT scanning doesn’t always provide the subtle variations on the endocast surface that are critical for correctly identifying convolutional details. The downside of course is possible damage to original specimens, and lack of sharing with colleagues at other institutions unless they visit the lab where these are made. Furthermore, the accuracy of virtual endocasts depends on the software, the researcher’s experience, expertise, and whether the algorithms used to correct for distortion, etc, are accurate.

What is the main current challenge in paleoneurology?

The major challenge is to synthesize the overall size data (ECV’s) with whatever sulcal and gyral information (e.g., lunate sulcus, fronto-orbital sulcus, Broca’s cap regions, etc) is available with morphometric analyses for each specimen with temporal and archaeological evidence, so that actual hypotheses can be generated than can be tested within (or even beyond) the paleoneurological community. This requires researchers fully cognizant of anatomical details, and both nonhuman primate and human neuroscience. Needless to say, but many more hominin and hominid endocasts need to be found and studied if paleoneurology is to become a better science.

Advices to those who begin working in this field …

Know your neuro- and cranial anatomies! Stay humble, lose your hubris, and keep in mind how rare endocasts are, and how imperfect these usually are, and how difficult, if not impossible it is to really know what the brain was like when the hominin was alive, and realize that you will probably never see an endocast that fully captures all the convolution details that were part of that once throbbing brain. A lack of hubris will be essential for good science, and don’t dismiss earlier works in paleoneurology simply because these are not modern or based on the last decade of morphometric advances. Staying up to date, or being current with the findings coming out of neuroscience will be particularly difficult.

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Australopithecus africanus

Amélie Beaudet and coauthors have published a shape analysis of the endocranial surface of two specimens included in the hypodigm of Australopithecus africanus, namely Sts 5 and Sts 60. They used a deformation-based approach, as to quantify the global endocranial surface variations. They compared the endocasts of the two specimens with modern humans (Homo sapiens) and chimps (Pan troglodytes and Pan paniscus). The endocranial geometry of the two piths is similar to the shape displayed by the apes. The incomplete Sts 60 lies right in between the two chimpanzees, while Sts 5 is slightly displaced toward the human form. The main difference between humans and the two australopiths can be localized in the parietal surface (larger in our species). Nonetheless, piths may present some minor shape difference on the frontal (orbital) cortex, when compared with chimps. Back in the ’20s Raymond Dart stated that, if extant African apes show a primitive scheme, a displacement of the lunate sulcus in Australopithecus might suggest a relative enlargement of the parietal cortex in these early hominids. Unfortunately, there are still disagreements on this point, because of the uncertainties associated with the identification of sulcal landmarks on fossil endocasts. Also frontal changes have been already proposed for other australopithecines. However, in this case the spatial interaction between frontal cortex and the facial system (orbits and ethmomaxillary block) may hamper a straight separation of the effects due to actual brain changes and those due to cranial constraints.

Precuneus and primates

The precuneus displays a remarkable variability in size and shape among adult humans, and it also represents a main difference between human and chimp brain morphology, being larger in our species. It can be argued that precuneus expansion in humans is due to an allometric pattern shared among primates. In this case, a large precuneus is a by-product of a big brain and scaling rules. We have now published a brain shape analysis in non-human primates, suggesting that this seems not the case. The midsagittal brain morphology in non-human primates is probably influenced by cranial architecture more than by brain differences. And, precuneus morphology is apparently not influenced by brain size, with no major differences between monkeys and apes. Therefore, its expansion in humans is likely to be a species-specific character, and not an allometric consequence of a large brain. The exact histological factors involved in this change is still to be investigated, as well as its functional (cognitive) consequences. In general, precuneus morphology is very variable also within other primate species, suggesting a noticeable plasticity. Its areas are crucial for coordination between body and vision (visuospatial integration, visual imaging, simulation, body cognition, autonoesis, etc.), and are influenced by both genetic and environmental factors (i.e., visuospatial training and practice). Its position physically matches those brain districts supposed to have undergone an expansion in the evolution of Homo sapiens, when compared with fossil hominids.

Brains and eyes

After our first survey on the morphological relationships between eyes and brains, here a comprehensive second study on this same topic. We have analyzed data from computed tomography (orbits and endocranial space) and magnetic resonance (eyes and brain), investigating modern humans, apes, and fossils. Soft tissue variation mainly deals with the distance between eyes and temporal lobes. Cranial variation mainly concerns the orientation of the orbits, probably influenced by parietal morphology and variation of the head functional axis. Phylogenetic differences are generally associated with the distance between orbits and braincase, with fossil humans showing an intermediate position between modern humans and apes. Here a Skull Box post with more details.

Sulcal imprints

The fuzzy geometry of the brain surface shapes the endocranial wall, and endocasts can show traces and imprints of the cortical sulcal patterns. Individual variation is noticeable, and the precise mechanisms behind these folding schemes are not clear at all. Hence, it is not recommended to use this information in a simplistic “phrenological” fashion, as unfortunately it has been done in many evolutionary studies. At the same time, cortical morphology is the direct result of neurons growth and development, and therefore even the pretentious rejection of this information seems unwise. Many authors dismiss any result based on brain gross morphology, simply because it is “just brain form”. This is probably because they ignore the developmental processes behind that forms, and they don’t take into account that when we talk about “brain form” we are implicitly referring to those processes, and not to a crude geometrical appearance. At least, sulcal patterns are useful (and the only available macroscopic) boundaries to detect the absolute or relative extension of some cerebral districts or cortical areas. So, despite all the uncertainties, they are directly providing information on cortical proportions. Proportions means “some areas are larger and some others are smaller”. Size is not always a matter of more or less neurons, but it is however matter of more or less “something”. Whatever it is, it should be functional, and maybe even adaptive some way, associated with some specific histological factor, or with some indirect physiological consequence. This is why the issue is not trivial.

Sulcal imprints are generally more visible on smaller and younger skulls. A recent study investigates the expression of the sulcal traces in macaques. Anterior folds (frontal and temporal lobes) leave more traces than the posterior ones (parietal and occipital). There are no many differences among young ontogenetic stages but then, during aging, the expression of the traces decreases noticeably, and imprints become more blurred. Local anatomical differences in the barrier between brain and skull (meninges, vessels, etc) can have a role in this size-related differences. Nonetheless, probably it is a matter of growth. In earlier ages, the brain generates a constant pressure on the vault bones, shaping the bone surface. But in later ages, when brain growth is concluded, that intimate physical relationship is looser. During aging, the brain even undergoes a shrinkage of about 7-8%, and the contact is further lost. This study is simple and effective, a good paper to approach the topic. Between an uncritical phrenological approach and a snobbish rejection of the evidence, we should consider an intermediate approach, in which we evaluate what kind of information we can obtain from these traits. To do that, we have to investigate their phenotypic factors and their mechanical influences, their structural associations and their variability.

Brainstorm …

As properly remarked in the prequel of the Planet of the Apes, we know everything about our brain, except how it does work. We are aware of such lack of knowledge, at least in theory. In practice, papers are replete of firm sentences and conclusive statements. But we use complex programs and devices, and we should not forget that these tools can only generate models of reality. Models based on algorithms that are trying to represent and simulate only some specific physical or spatial properties. Our brain models are but statistical outputs, not real “brains”. We identify brain activity through indirect blood or metabolic functions, assuming there is a strict correspondence between those signals and our concept of “at work”. A correspondence that is reasonable, but not that strict. Even basic anatomical issues can be blurred after a more detailed scrutiny, mostly when previous knowledge is based on information that has been copy-and-pasted through decades. We are more and more finding strange factors influencing our results. Apparently, the brain undergoes daily variations, and the braincase may suffer seasonal changes. Brain structure and function can be even influenced by head position and posture. These unexpected effects recommend further caution when making too general conclusions from specific and punctual results. Let’s take into account that we still miss much information on gross neuroanatomical components. For example, we still ignore the function of the cerebellum, that has four time the number of neurons of the brain, and we still don’t know all the functions of the glial cells, that may be nine times more numerous than neurons. And, we don’t know how much brain anatomy and functions are the result of genetic programs or environmental influences. In only few weeks, training can easily improve or demote brain complexity. Nothing new under the sun: science is about hypotheses, and hypotheses need to be tested and validated. Our models are tentatively designed with this scope in mind. This summer post is a summary of articles concerning some methodological limitations and some curious result dealing with brain structure and function. And an invitation to interpret results for what they are: evidences supporting or rejecting hypotheses. Remember that those are not neurons: just pixels! Take it easy …

Language and fossils

This week I have published an opinion paper on language and paleoneurology, in a volume of Frontiers in Human Neuroscience dedicated to language, skulls, and brains. I review the fossil evidence on language, suggesting that most of such evidence concerns brain areas that are influenced by cranial structural constraints, or is based on speculations associated with individual bone remains. Thus, strictly speaking, there is no consistent evidence on language evolution when you deal with fossil anatomy. Ralph Holloway already stressed this point before, but it seems that most books and articles introducing this topic simply keep on stating the opposite, following a mantra (usually void of citations) according to which fossils must clearly reveal the cerebral (usually frontal) changes behind language evolution. The lack of scientific evidence in this context does not mean that there is no association between language and brain evolutionary changes in hominids, but just that fossils can provide only a very incomplete (and insufficient) view of this process. Firm statements, scientifically speaking, should be avoided, and relegated to storytelling and science marketing.

The dorsal and medial parietal areas, probably larger in Neanderthals and even more derived in modern humans, are not generally considered when discussing language processes, and most of the debate has been centred on frontal lobe functions. Nonetheless, the parietal areas are crucial for hand coordination and manipulative abilities, both factors that have always been regarded as influential in language evolution. Also, recent evidence suggests that language has an important embodied component: language coding passes through body experience and simulation, something which is profoundly associated with the functions of the deep parietal folds. Therefore, we should consider whether changes in the whole fronto-parietal system may have triggered or facilitated language in the human genus. The paper is open access.


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