For the first time in many years, I’m offering a new advanced undergrad seminar here at Vassar. When I arrived here 8 years ago, I was mainly thinking about Homo naledi and ontogeny, so those were the foci of my seminars. But my research has begun looking more at brain evolution and especially the evidence from fossil endocasts, and there is a lot of literature I need to catch up on.
So I’ve invited students along for this brainstorm, using the question “Is the human brain special?” as a starting point to learn about how the beautifully congealed soup sloshing around inside our skull makes us such quirky animals. In the first half of the semester we’ll read up on brain anatomy and structure, and students will use some of the fossil endocast data I’ve accrued over the years to learn more about a given brain region and extinct hominin. In the second half of the semester we’ll read about the brains, behavior, and endocast fossils of very distant relatives — invertebrates, birds, whales, and dogs — that have been celebrated for their own ‘advanced intelligence.’ We’ll also read about how the evolution of our brains may have predisposed us to certain conditions like addiction and Alzheimer’s, and how brain science has been exploited toward racist and sexist ends (increasingly relevant in America today, sadly).
It will be a lot of work (I’m a very slow, distractible reader) but I’m excited to delve into this literature and see what insights our super sharp students here at Vassar come up with in discussions and projects. The course syllabus (ANTH 323) is available on my Teaching page — I’d be keen to hear suggestions for readings and assignments from folks who know more about brains than I do!
In the Summer of 2019 I worked with some great Vassar undergrads to make virtual endocasts and generate new brain size estimates for the Neandertals from the site of Krapina, which we then published in 2021 (discussed in this blog post). The virtual approach to endocast reconstruction uses 3D landmark-based geometric morphometrics methods, and so in the spirit of open science we also published all the landmark data used for the study (as well as a bunch of other fossil human brain size estimates) in the Zenodo repository (here).
Neandertal fossil specimens Krapina 3 (purple/green) and Krapina 6 (yellow/red) with preserved landmarks and virtually reconstructed endocasts.
Something major and global happened around that time — who can even remember what? — and so I never got around to posting R code to accompany the study. So, I’ve finally gotten around to adding some very basic code to the Zenodo entry (better late than never). The code simply reads in the landmarks, estimates missing data for fossils, and does some very basic shape analysis and visualization. It’s doesn’t get into all the nuts and bolts of our study, but it should be enough to help folks check our data or get started with shape analysis in R.
R code includes ways to visualize the landmark data. Left: Principal components analysis graph of endocast shape for humans (red) and Neandertals (blue). Right: Triangle meshes of the average human and Neandertal endocast shapes, viewed from the right, bottom, and back.
Original article Cofran Z, Boone M, Petticord M. 2021. Virtually estimated endocranial volumes of the Krapina Neandertals. American Journal of Physical Anthropology 174: 117–128. (link)
What makes the human brain special, and how did it change throughout our evolutionary history? One way to answer this question by comparing actual brains or MRI scans of living animals. But only fossils can show what changed and when over the past several million years, and sadly brains are basically an elaborately congealed soup that doesn’t stay fresh upon death, so they never fossilize (well, almost never). Happily, though, bones can preserve for millions of years, and they are literally molded by their soft and squishy surroundings. As the brain grows, it pushes outward against the inner surface of the skull, which can save the scars of the submerged cerebrum: nerds like me call these impressions an “endocast.”
Endocasts of Homo naledi (pink) and Homo erectus (yellow). Fossils are viewed from the left side and are variably preserved.
Nicole Labra and Antoine Balzeau have led a cool study, hot off the press, examining what such endocasts can tell us about the underlying brain anatomy. Importantly, they show how difficult it is to clearly and consistently identify many brainy boundaries. This is very salient in “paleoneurology,” the study of brain evolution especially based off endocasts: the problem probably best illustrated by the nearly century-long debate about the natural endcoast of the “Taung child” fossil (Australopithecus africanus).
Labra & colleagues used a clever approach to address this paleontological and epistemological problem. They first generated an endocast directly associated with its brain from an MRI scan of a living human, allowing them see precisely where specific brain grooves (“sulci”) lay relative to the endocast surface. They then asked a bunch of researchers—myself included—to try to identify sulci on the endocast, and then looked at how our responses compared to both one another’s and to the actual, known sulcus positions.
Figure 1 from Labra et al. (in press) showing how the brain and endocast were obtained and analyzed.
Their analysis showed that we varied quite a bit in our identifications on the endocast. As Emiliano Bruner (who also participated) discusses in his blog post, we tended to identify the stronger impressions toward the bottom and sides of the endocast better and more consistently. Some of this variability and uncertainty among researchers is due to the faintness and incompleteness of many brain impressions, and some due to biased expectations about where a given sulcus “should” be based on our previous experiences and published references.
When Antoine Balzeau first contacted me about this project, I was just beginning to dabble in paleoneurology, learning some brain anatomy for the first time for a description of an old Australopithecus endocast called “MLD 3.” I initially thought MLD 3 would be a quick and simple study—boy was I spectacularly disappointed!
Figure 3 from Cofran et al. 2023, comparing two different chimpanzee brains, and two corresponding interpretations of the MLD 3 endocast.
Probably reflecting observer bias and desire for definitive results, we initially interpreted the endocast impressions on MLD 3 as representing a ‘human-like’ anatomy that is super rare in living chimpanzees (namely the “LS” depicted in the right half of the figure above). The researchers who peer-reviewed the first draft of our paper, though, suggested we be more cautious in our interpretations; one reviewer outright disagreed with us in support of a more ‘ape-like’ interpretation (left half of the figure above). The review process alone underscored the subjectivity and uncertainty in analyzing endocasts. In the end we presented both interpretations, and I honestly don’t know which (if either) is most likely to be correct. So the study by Labra and colleagues provides a nice empirical illustration of this cranial conundrum.
Fortunately, researchers are developing methods to help identify brain structures on endocasts. Amélie Beaudet, Jean Dumoncel, and Edwin de Jager among others have done some really impressive work looking at variability in both brains (for instance here) and endocasts (for instance here). By using computer-based 3D data and methods, these researchers have shown where many brain sulci tend to be located (see here). By developing a better understanding of variation in where sulci sit on an endocast, we can have a better idea of which sulci might be represented on fossil endocasts, which in turn can tell us about the brains of our extinct relatives. Edwin and Amélie presented a very cool new analysis of Australopithecus/Paranthropus boisei endocasts, building off this digital approach, at the recent ESHE conference. And as noted in our MLD 3 paper, I think machine learning and other ‘artificial intelligence’ approaches could also help us identify ambiguous features from frustrating fossil fragments.
Each year in my intro bio-anthro class, we start the course by asking how our brains contribute to making us humans such quirky animals. Our first lab assignment in the class uses 3D models of brain endocasts, to ask whether modern human and fossil hominin brains are merely primate brains scaled up to a larger size. In the Before Times, students downloaded 3D meshes that I had made, and study and measure them with the open-source software Meshlab. But since the pandemic has forced everyone onto their own personal computers, I made the activity all online, to minimize issues arising from unequal access to computing resources. And since it’s all online, I may as well make it available to everyone in case it’s useful for other people’s teaching.
The lab involves taking measurements on 3D models on Sketchfab using their handy measurement tool, and entering the data into a Google Sheets table, which then automatically creates graphs, examines the scaling relationship between brain size (endocranial volume, ECV) and endocast measurements, and makes predictions about humans and fossil hominins based off the primate scaling relationship. Here’s the quick walk-through:
Go to the “Data sources” tab in the Google Sheet, follow the link to the Sketchfab Measurement Tool, and copy the link to the endocast you want to study (3D models can only be accessed with the specific links).
Following the endocast Sketchfab link (column D) will bring you to a page with the 3D endocast, as well as some information about how the endocast was created and includes its overall brain size (ECV in cubic cm). Pasting the link when prompted in the Measurement Tool page will allow you to load, view, and take linear measurements on the endocast.
Hylobates lar endocast, measuring cerebral hemisphere length between the green and red dots.
Sketchfab makes it quite easy to take simple linear measurements, by simply clicking where you want to place the start and end points. The 3D models of the endocasts are all properly scaled, and so all measurements that appear in the window are in millimeters.
The assignment specifies three simple measurements for students to take on each endocast (length, width, and height). In addition, students get to propose a measurement for the size of the prefrontal cortex, since our accompanying reading (Schoenemann, 2006) explains that it is debated whether the human prefrontal is disproportionately enlarged. All measurements are then entered into the Google Sheet — I wanted students to manually enter the ECV for each endocast, to help them appreciate the overall brain size differences in this virtual dataset (size and scale are often lost when you have to look at everything on the same-sized 2D screen).
Feel free to use or adapt this assignment for your own classes. The assignment instructions can be found here, and the data recording sheet (with links to endocast 3D models) can be found here — these are Google documents that are visible, but you can save and edit them by either downloading them or making a copy to open in Docs or Sheets.
In the latest paper out of the lab (here), my students and I reconstructed the brain endocasts of the Krapina Neandertals. The Krapina rock shelter in Croatia is a remarkable site. Dating to around 130,000 years ago (if not older), the Krapina fossils are early members of the Neandertal lineage. In addition, the fossils represent dozens of Neandertals, from infants to adults. Part of what drew me to the site were the juvenile skulls, since they can tell us about growth and development in these early humans. But, the fossils are quite fragmentary, and needed to be reconstructed to estimate important characteristics like brain size.
Figure 1 from our paper, showing the five Krapina crania (A & B are the same individual) with the endocranial surface highlighted.
Vassar College has a great program called URSI, where students team up with faculty to get hand on experience conducting research over the summer. So, two summers ago my students and I worked on virtually putting these Humpties Dumpty back together again. Using 3D surface scans of the original fossils and CT scans of modern humans, we used virtual methods to digitally reconstruct the endocasts, which are a good proxy for brain size and shape. Here’s the basic workflow:
Figure 2 from the paper, depicting the workflow for virtually reconstructing fossil endocasts, represented by the famous Krapina 3 or “C” cranium.
The human endocasts were produced from recent humans from the Terry Anatomical Collection, generously made available here by Dr. Lynn Copes. We have posted the 3D landmark data for the humans, the preserved landmarks from the Neandertals, and a big list of estimated brain sizes for Neandertals, in the open access repository Zenodo (here). So, hopefully anyone can repeat our results, or use these data in their own research.
With virtual methods, we could generate multiple reconstructions of each Neandertal fairly easily, giving an idea of how certain or uncertain our brain size estimates were. In the end, we showed that i) the Krapina juveniles, who were probably around 6-7 years old, had brain sizes within the adult range (it’s same with modern humans); ii) average brain size at Krapina was a little lower than previously estimated; and iii) although later Neandertals from other sites had larger brains on average, the difference is not necessarily greater than could be expected by chance.
I’ve participated in Vassar’s URSI program for the past few years and it has been a lot of fun. Last (virtual) summer, my students and I compared hip growth in humans and Australopithecusafricanus, and this coming summer we will examine the brains of the greatest animals of all time — gibbons!
Lawnchair Anthropology 