A group coordinated by Dr. Vera Weisbecker examined whether the evolution of mammalian brain partitions follows conserved developmental constraints, causing the brain to evolve as an integrated unit in which the partitions scale with brain size. According to this ‘late equals large’ hypothesis, the timing of neurogenesis predicts the size of the partition such that later and more extended neurogenesis produces larger partitions due to the production of more neural precursors. In order to investigate the impact of neurogenesis on patterns of brain partition growth, the volumes of the whole brain and major partitions were reconstructed from soft-tissue diceCT scans of three marsupial species, including individuals with ages ranging from 1 day to adulthood. They tested three hypotheses consistent with a conserved brain partition growth: H1 postulates that partition scaling during development reflects the evolutionary partition scaling, and thus growth patterns should be uniform between species; H2 assumes that a neurogenesis-driven pattern of partition scaling is predictable from adult brain size, i.e. brain partitions scale with brain size; and H3 states that growth with age might differ between species according to brain size and/or neurogenetic events. Regressions of log partition volume against log rest-of-the-brain volume (whole-brain volume minus partition volume) showed significant interspecific differences in slopes and intercepts of most brain partitions, indicating diverse scaling patterns between species, which could not be predicted by adult brain size, as the smallest-brained species had intermediate slope to the other two. Growth curves of log partition volume against age were similar in all partitions within-species, but differed between species, particularly in growth rates, with the species with intermediate brain size having slower rates than the other two. Differences in growth patterns do not seem to be related to neurogenetic schedule as largest partitions are not especially late in their development and important maturation processes, like eye opening, occur closer to the end of the growth phase. Thus, none of the hypotheses are supported by these results, challenging the conserved neurogenetic schedules behind the evolution of mammal brain partitions. Moreover, the authors found high phylogenetic signal in brain partition scaling, revealing that a large part of the scaling relationship between brain and partition volumes is explained by phylogeny, which is more in agreement with a mosaic evolution of brain partition sizes, stressing its biological meaning and the level of mammalian brain plasticity. However, the intraspecific regular partition growth curves led the authors to contemplate the existence of an early brain partition pattern regulated by regional gene expression, and propose that further studies of brain partition evolution should integrate developmental neuromere expression models, neuron density, and patterns of neuron migration.
The mature primate brain consists of many layers with the outer layer or cerebral cortex forming folds known as sulci and gyri. During embryonic development, the brain is divided into zones with the inner-most ventricular zone where neurons are formed and a series of cytoarchitecturally distinct layers forming plates radiating outward. The subplate is located between the inner ventricular zone and the outer cortical plate hosting the migration of neurons allowing brain expansion. Most embryonic brain research is conducted on non-primate mammals but there are substantial differences in the development of the non-primate and primate brain. A very recent study utilized existing primate tissue databases to examine the embryonic development of the subplate zone in non-human and human primates. Duque et al. found during that development of the macaque brain, once the neurons have migrated to the subplate they then are pushed downward by axons derived from the subcortical layer before further compression occurs from further axonal development originating from the cortical layer. The implications of this force acting on the neurons within the subplate suggests that thickness of the subplate differs unevenly throughout the brain potentially due to an increased axonal density. Duque et al. suggest the density of axonal fibers increases with demand for more connectivity between brain regions with those areas possessing a high-demand for greater complexity causing a thicker subplate.
Changes at the cellular-level of the subplate also have implications for the development of the cerebral convolutions such as sulci and gyri. It was recently posed that the folding patterns in the human brain are the result of mechanical forces related to the subplate and outer expansion of the cerebral cortex. Tallinen et al. showed through numeric and physical simulations with the support of MRI that during fetal development the subplate stabilizes while the outer cortical plate continues to expand. The final stages of growth see the cortical layer undergo extensive gyrification to form the folding patterns we see in the adult human brain. Overall, a better understanding of human neurobiology informed through non-human primate neurobiology offers a glimpse into the evolutionary pathways which led to the evolution of modern humans.
In their last review Jean-Jacques Hublin, Simon Neubauer and Philipp Gunz address the effects of hominins’ life histories in brain evolution. Encephalization in humans involved energetic costs that were sustained through changes in social structure and metabolic adaptations, including changes in the diet quality, as explained by the Expensive Tissue Hypothesis, and the ability to store energy in fat tissue, the primary source of energy for neonate brains. Because of the constraints of birth-giving associated with bipedalism, human brains develop mainly after birth. Also because of this prolonged development, the brain is exposed to a rich environment during its wiring process, with the child furthermore protected by the social community.
The study of fossil hominins’ brain development is only possible through the analysis of their endocranial casts, using cross-sectional samples. To establish their life histories it is necessary to attribute an age at death, generally according to the eruption timing of the teeth. Beyond variation in brain size, signs of brain reorganization can be investigated in fossil species to get insights about their cognitive capacities. Regarding Australopithecus, it is not yet clear how their brain development and life histories were more like that of humans or that of chimps. Homo erectus had body proportions and social structures closer to that of modern humans, but different brain organization and smaller brain capacity than that of latter Homo, pointing to a faster life history. Homo sapiens and H. neanderthalensis separately evolved similar brain capacities, although with different morphologies, and had different life histories, faster in the latter. Brain organization differs soon after birth, as modern humans undergo the so-called globularization phase, which does not exist in chimps or Neanderthals. This globular shape of the brain is characterized by the bulging of the parietal areas which may be linked to reorganization of the internal regions, like the precuneus.
There is still a debate about whether or not life histories in fossils can be investigated in terms of brain growth and development. The different brain morphology of Australopithecus when compared with African apes suggests that brain reorganization could have pre-dated encephalization. At last, our patterns of socio-cultural evolution might have been fundamentally responsible for the adaptive changes essential for the evolution of our big brains.