Pre-publication: Brain growth in Homo erectus (plus free code!)

The annual meetings of the American Association of Physical Anthropologists were going on all last week, and I gave my first talk before the Association (co-authored with Jeremy DeSilva). The talk focused on using resampling methods and the abysmal human fossil record to assess whether human-like brain size growth rates were present in our >1 mya ancestor Homo erectus. This is something I’ve actually been sitting on for a while, and wanted to wait til after the talk to post for all to see. I haven’t written this up yet for publication, but before then I’d like to briefly share the results here.

Background: Humans’ large brains are critical for giving us our unique capabilities such as language and culture. We achieve these large (both absolutely, and relative to our body size) brains by having really high brain growth rates across several years; most notable are exceptionally high, “fetal-like” rates during the first 1-2 years of life. Thus, rapid brain growth shortly after birth is a key aspect of human uniqueness – but how ancient is this strategy?

Materials: We can plot brain size at birth in humans and chimpanzees (our closest living relatives) to visualize what makes humans stand out (Figure 1).

Figure 1. Brain size (volume) at given ages. Humans=black, chimpanzees=red. Ranges of brain size at birth, and the chronological age of the Mojokerto fossil, in blue.

Human data come from Cogueugniot and Hublin (2012), and chimpanzees from Herndon et al. (1999) and Neubauer et al. (2012). The earliest fossil evidence able to address this question comes from Homo erectus. Because of the tight relationship between newborn and adult brain size (DeSilva and Lesnik 2008), we can use adult Homo erectus brain volumes (n=10, mean = 916.5 cm^3) to predict that of the species’ newborns: mean = 288.9 cm^3, sd = 17.1). An almost-recent analysis of the Mojokerto Homo erectus infant calvaria suggests a size of 663 cm^3 and an age of 0.5-1.25 years (Coqueugniot et al. 2004; this study actually suggests an oldest age of 1.5 years, but the chimpanzee sample here requires us to limit the study to no more than 1.25 years). Because we have a H. erectus fossil less than 2 years of age, and we can estimate brain size at birth, we can indirectly assess early brain growth in this species.

Methods: Resampling statistics allow inferences about brain growth rates in this extinct species, incorporating the uncertainty in both brain size at birth, and in the chronological age of the Mojokerto fossil. We thus ask of each species, what growth rates are necessary to grow one of the newborn brain sizes to any infant between 0.5-1.25 years? And from there, we compare these resampled growth rates (or rather, ‘pseudo-velocities’) between species – is H. erectus more similar to modern humans or chimpanzees? There are 294 unique newborn-infant comparisons for humans and 240 for the chimpanzee sample. We therefore compare these empirical newborn-infant pairs from extant species to 7500 resampled H. erectus pairs, randomly selecting a newborn H. erectus size based on the parameters above, and randomly selecting an age from 0.5-1.25 years for the Mojokerto specimen. This procedure is used to compare both absolute size change (the difference between an infant and a newborn size, in cm^3/year), and and proportional size change (infant/newborn size).

Results: Humans’ high early brain growth rates after birth are reflected in the ‘pseudovelocity curve’ (Figure 2). Chimps have a similar pattern of faster rates earlier on, but these are ultimately lower than humans’. Using the Mojokerto infant’s brain size (and it’s probable ages) and the likely range of H. erectus neonatal brain sizes (mean = 288, sd = 17), it is fairly clear that H. erectus achieved its infant brain size with high, human-like rates in brain volume increase.

Figure 2. Brain size growth rates (‘pseudo-velocity’) at given ages. Humans=black, chimpanzees=red, and Homo erectus,=blue.

However, if we look at proportional size change, the factor by which brain size increases from birth to a given age, we see a great deal of overlap both between age groups within a species, and between different species. Cross-sectional data create a great deal of overlap in implied proportional size change between ages within a species; it is easier to consider proportional size change between taxa, conflating ages, then  (Figure 3). Humans show a massive amount of variation in potential growth rates from birth to 0.5-1.25 years, and chimpanzees also show a great deal of variation, albeit generally lower than in the human sample. Relative growth rates in Homo erectus are intermediate between the two extant species.

Figure 3. Proportional brain size increase (infant/newborn size). 

Significance: Brain size growth shortly after birth is critical for humans’ adaptative strategy: growing a large brain requires a lot of energy and parental (especially maternal) investment (Leigh 2004). Plus, in humans this rapid increase may correspond with the creation of innumerable white-matter connections between regions of the brain (Sakai et al. 2012), important for cognition or intelligence. The H. erectus fossil record (1 infant and 10 adults) provides a limited view into this developmental period. However, comparative data on extant animals (e.g. brain sizes from birth to adulthood), coupled with resampling statistics, allow inferences to be made about brain growth rates in H. erectus over 1 million years ago.

Assuming the Mojokerto H. erectus infant is accurately aged (Coqueugniot et al. 2004), and that Homo erectus followed the same neonatal-adult scaling relationship as other apes and monkeys (DeSilva and Lesnik 2008), it is likely that H. erectus had human-like rates of absolute brain size growth. Thus, the energetic and parental requirements to raise such brainy babies, seen in modern humans, may have been present in Homo erectus some 1.5 million years ago or so. This may also imply rapid white-matter proliferation (i.e. neural connections) in this species, suggesting an intellectually (i.e. socially or linguistically) stimulating infancy and childhood in this species. At the same time, relative brain size growth appears to scale with overall brain size: larger brains require proportionally higher growth rates. This is in line with studies suggesting that in many ways, the human brain is a scaled-up version of other primates’ (e.g. Herculano-Houzel 2012).
This study was made possible with published data, and the free statistical programming language R.

Contact me if you want the R code used for this analysis, I’m glad to share it!!!

Coqueugniot H, Hublin JJ, Veillon F, Houët F, & Jacob T (2004). Early brain growth in Homo erectus and implications for cognitive ability. Nature, 431 (7006), 299-302 PMID: 15372030

Coqueugniot H, & Hublin JJ (2012). Age-related changes of digital endocranial volume during human ontogeny: results from an osteological reference collection. American journal of physical anthropology, 147 (2), 312-8 PMID: 22190338

DeSilva JM, & Lesnik JJ (2008). Brain size at birth throughout human evolution: a new method for estimating neonatal brain size in hominins. Journal of human evolution, 55 (6), 1064-74 PMID: 18789811

Herculano-Houzel S (2012). The remarkable, yet not extraordinary, human brain as a scaled-up primate brain and its associated cost. Proceedings of the National Academy of Sciences of the United States of America, 109 Suppl 1, 10661-8 PMID: 22723358

Herndon JG, Tigges J, Anderson DC, Klumpp SA, & McClure HM (1999). Brain weight throughout the life span of the chimpanzee. The Journal of comparative neurology, 409 (4), 567-72 PMID: 10376740

Leigh SR (2004). Brain growth, life history, and cognition in primate and human evolution. American journal of primatology, 62 (3), 139-64 PMID: 15027089

Neubauer, S., Gunz, P., Schwarz, U., Hublin, J., & Boesch, C. (2012). Brief communication: Endocranial volumes in an ontogenetic sample of chimpanzees from the taï forest national park, ivory coast American Journal of Physical Anthropology, 147 (2), 319-325 DOI: 10.1002/ajpa.21641

Sakai T, Matsui M, Mikami A, Malkova L, Hamada Y, Tomonaga M, Suzuki J, Tanaka M, Miyabe-Nishiwaki T, Makishima H, Nakatsukasa M, & Matsuzawa T (2012). Developmental patterns of chimpanzee cerebral tissues provide important clues for understanding the remarkable enlargement of the human brain. Proceedings. Biological sciences / The Royal Society, 280 (1753) PMID: 23256194


Fun with Einstein’s Brain

I stumbled across a little blurb today in ScienceNow about a new study by Dean Falk (University of Michigan PhD, 1976!) about Einstein’s brain. Luckily, back in good olde 1955, when urban money was moving to suburbs and Marty McFly was trying not to screw up the future, people realized that the recently-late Einstein was a genius whose brain needed to be preserved. Fun facts that I found out:

Einstein’s brain was only 1230 cc. The average for modern people is around 1400 (Holloway 2000). Here’s a list of ancient Homo fossils, from Holloway (2000) and some others, whose cranial capacities are about the same as or greater than Einstein’s:

Zhoukoudian X (Chinese H. erectus, ~1225 cc), Ngandong 10 (Javanese H. erectus, ~1231 cc); Kabwe, LH 18, Eyasi, Saldanha, BOU-VP 16/1 (African “archaic” Homo sapiens); Narmada, Jinniushan, Yinkou (Asian “archaic” Homo sapiens); Vertesszolos 2, Reilingen, Steinheim, Swanscombge, Fontachevade, Ehringsdorf, Biache, Petralona, Atapuerca 4 (European “archaic” Homo sapiens); and most Neandertals.

This shows that, while brain size was important in the evolution of human cognition, it is not everything. I mean, how many of these hominins could begin to fathom something like special relativity? Of course, back in the Paleolithic, when life was hard and one has to worry about how to obtain food, ward off predators and persist in some sort of society, who had time for such things? On the other hand, I’m a modern human–I have no idea how large my brain is–but I can barely wrap my mind around most things in physics. So it seems that human cognition–even genius-level, such as Einstein’s–is founded in biology, but also culture and environment.

The article also suggests to me that no one really knows how the brain works. Yes, the parietal regions are associated with maths and such, and Einstein had relatively large parietal lobes. But how and why do one person’s parietal lobes confer greater math capabilities than another? (If the parietal lobes relate to mathematical ability, I might lack these)

The article also tells that Falk found a “knob-like structure” in the motor cortex, and that such “knobs” have also been associated with musical abilities. I’m not a neuroscientist, and I don’t know what these ‘knobs’ are. But it sounds like scientists kind of know what these do, since they see these structures more in people who are notable for a given talent (math, music, etc) But are these inherent in the brain and allow people these special abilities, or are they more environmental in origin, arising from certain experiences and exposures? More importantly, what do these do?! Falk also found other brain abnormalities, “that she speculates might somehow be related to Einstein’s superior ability to conceptualize physics problems.” This may well be the case, but it is still unclear why this should be so.

So I think this study is great, because it can provide neuroscientists with bases for future research on brain function and anatomy. At the same time, it underscores the fact that as smart as we humans are, we don’t yet understand how or why we are so special.


Holloway, R.L. 2000. Brain. In: Delson et al, eds. Encyclopedia of Human Evolution and Prehistory. New York: Garland Publishing, Inc. p 141-149.

Guest Post: Jerry and Julie Jive

“Good news, everyone!” to quote Prof. Farnsworth. Our good friends Jerry DeSilva and Julie Lesnik just published a paper in the Journal of Human Evolution, about neonatal brain size in primates [1]. Rather than talk and talk about it, probably missing the important stuff, I made some calls. The authors were kind enough to make a cameo appearance at Lawn Chair to talk about their paper about their paper. Thanks, Jerry and Julie! Here’s what they had to say:


This paper presents a regression equation that can be used to calculate the size of the brain at birth in different hominin species.


Knowing the size of the brain at birth is critical for understanding obstetric constraints and brain development throughout human evolution. Unfortunately, it is very unlikely to find fossil evidence of how big the brain was at birth in human ancestors (though see below). This paper presents a way to get around the absence of fossil evidence and calculate the size of the neonatal brain in early homs using what we know about brain development in modern primates.

Things Jerry liked about the paper:

Humans are so unusual, and in biological anthropology we often study ways in which humans are different from other primates. However, what this study finds is that humans are no different from other primates in terms of the adult/neonatal brain scaling relationship. This means that we have exactly the brain size at birth expected given the size of our brains as adults. Because of this, we can infer that our extinct ancestors and relatives also followed this ‘rule’ of adult/neonatal brain size, and can calculate the size of the brain at birth from reliable estimates of brain size in 89 adult fossil crania that have been unearthed.

I am also thrilled that Julie and I may have solved the “% brain size at birth” issue that has been all over the literature lately. Did Homo erectus have a more human-like or a more chimpanzee-like pattern of brain growth? What about australopiths? Well, we’ve found that the whole issue of % brain size at birth is simply a function of the scaling relationship between adult and neonatal brain size. Because they do not scale 1:1, but instead scale 1:0.73 (roughly), as the adult brain gets bigger, the neonatal brain gets proportionately smaller. Therefore, less of brain growth occurs in the womb as overall adult brain size increases. If you know the size of a hominin brain as an adult (which we do from the many preserved fossil crania), you can calculate the size of the brain as a baby, and then easily take a % of how much of that brain growth is achieved by birth.

Again, because of the negative allometry (m=0.73), we argue that % of brain size at birth in hominins was never “chimpanzee-like” or “human-like”, but instead followed a gradual progression from a chimpanzee-like ancestral condition to what we have today.

Things Julie liked about the paper:

So much is going on when we think about hominid evolution, especially in the early Pleistocene. With the emergence of Homo brain size is increasing, bipedality is becoming more efficient, and tool use is becoming more advanced. What I like about this paper is that understanding neonatal brain size is one way of tying all of those elements together. Humans are considered to be secondarily altricial meaning that they are born in a more underdeveloped state than their ancestors. Selection for this smaller neonatal size is often considered to be linked to the constraints placed on the pelvis by selection for more efficient bipedal locomotion. A small brain size at birth and a large adult brain always seemed exceptional for Homo. What our paper shows is that the relationship is entirely normal across anthropoids. So, where is the selective pressure? On the larger brain as an adult or on the smaller brain as a newborn? I am now more apt to lean towards larger adult brain. Efficient bipedality is important for exactly that reason; it’s efficient and therefore requires less energy to walk upright and allows the body to allot that energy to other tasks, such as maintenance of a large brain. Add tool-use advancement to the equation and it seems bigger brains and more advanced cognitive abilities were of primary importance at this stage of human evolution.

What we’d do different:

I would have included Neandertals. Julie and I made a statement in the introduction that the discovery of neonatal crania was bordering on impossible. Just days before our paper appeared on-line, however, Marcia Ponce de Leon published a fantastic paper in PNAS on a neonatal Neandertal cranium from Mezmaiskaya Cave in Russia [2]. What is very exciting to me is that this newly described fossil allows us to test our regression equation. How accurate is it in predicting the size of the brain at birth in Neandertals (which we now know because of this new specimen)? Our regression would predict a brain size of about 425 cc, which is very close to the size of the brain at birth in the Mezmaiskaya infant and well within the 95% CI. When two independent methods arrive at the same result, it is reasonable to argue that the method is valid.


1. DeSilva J, and Lesnik J. 2008. Brain size at birth throughout human evolution: a new method for estimating neonatal brain size in hominins. Journal of Human Evolution, corrected proof in press.

2. Ponce de Leon M, Golovanova L, Doronichev V, Ramanova G, Akazawa T, Kondo O, Ishima H, and Zollikofer C. 2008. Neanderthal brain size at birth provides insights into the evolution of human life history. Proceedings of the National Academy of Sciences 105: 13764-13768