Pattern of Gyrification

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Human Uniqueness Compared to "Great Apes": 
Likely Difference
Human Universality: 
Individual Universal (All Individuals Everywhere)
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Attempts have been made to discriminate changes in neurological reorganization within the hominin lineage by comparing the endocasts, real or virtual molds of the cranial case the brain sits within, of extinct hominids. Two areas are of particular interest in humans: 1) the presence of a ramified sylvian fissure which delimit Broca’s area (an area known to be important in speech formation), which replaces a unique anatomically consistent fissure in apes; and 2) the loss of the lunate sulcus, delimiting the primary visual cortex, which is otherwise found in living apes. Endocast studies of each of these regions are difficult to interpret as they are contingent upon impressions left on the interior of the skull by the brain during development; further while some gross similarities can be drawn across a species there is still a large amount of variation in individuals. The latter is confounded by limited numbers of examples of semi-intact skulls from given hominid species as well as difficulties in measurement which remain highly controversial (where different individuals from the same species, recorded by different researchers, have given opposing results). What appears to be true is that most hominids, since Australopithecus, seem to have both of these features. However, the fossil record remains spotty and inconclusive for some species and complete conclusions cannot yet be drawn. The existence of another related feature, the petalias (an asymmetry between the left and right lobes) suggests a divergence event sometime during the brain expansion after H. habilis.

Background Information: 

Gyrification (or cortical folding) is the the process by which the brain’s surface forms ridges and valleys (e.g. gyri and sulci). This occurs during neuronal migration over the course of development. As the layers of cortex are established, interconnections grow between neurons. Gyri are thought to be formed by the tensile strength of these neuronal connections. Thus, areas that are densely interconnected will pull together to form gyri whereas areas loosely connected will drift apart and form sulci. Bringing densely interconnected regions closer together also functions to promote efficient processing, as there is less distance to travel between closely related neurons. In humans, the greatest development of gyrification occurs during the third trimester of pregnancy, when the relatively smooth (lissencephalic) brain starts to contort into its adult, wrinkled morphology. The overall degree of cortical folding can be quantified by the gyrification index.
Increased gyrification presents an alternative way to increase the amount of neurons packed into the brain without increasing the overall volume. That is, gyrification allows more cortical surface area to be packed into the same amount of space than is possible with a lissencephalic brain. In fact, the human cortical sheet is estimated to have about three times larger area as the inner surface of the skull. The greater amount of cells and efficiency in processing is hypothesized to allow for greater cognitive abilities. For instance, the degree of folding in temporo-occipital cortex is positively correlated with intelligence scores in humans, and many mental disorders are associated with lissencephaly and the absence of normal gyri and sulci. However, the relationship between gyrification and overall neuron number is complex - greater overall neuron count does not necessarily equal greater cognitive capacity. Indeed, some mental disorders such as autism are associated with greater cell number and gyrification due to insufficient neuronal pruning. This is further complicated by the occurrence of autistic savants, individuals with severe deficits in some cognitive domains (typically including social deficits) at the same time as incredible abilities in other cognitive domains (typically in mathematics and declarative memory).

The Human Difference: 

Attempts have been made to discriminate changes in neurological reorganization within the hominin lineage by comparing the endocasts, real or virtual molds of the cranial case the brain sits within, of extinct hominids. Two areas are of particular interest in humans: 1) the presence of a ramified sylvian fissue which delimit Broca’s area (an area known to be important in speech formation), which replaces a unique anatomically consistent fissure in apes; and 2) the loss of the lunate sulcus, delimiting the primary visual cortex, which is otherwise found in living apes. Endocast studies of each of these regions are difficult to interpret as they are contingent upon impressions left on the interior of the skull by the brain during development; further while some gross similaries can be drawn across a species there is still a large amount of variation in individuals. The latter is confounded by limited numbers of examples of semi-intact skulls from given homid species as well as difficulties in measurement which remain highly controversial (where different individuals from the same species, recorded by different researchers, have given opposing results). What appears to be true is that most hominids, since Australopithecus, seem to have both of these features. However, the fossil record remains spotty and inconclusive for some species and complete conclusions cannot yet be drawn. The existence of another related feature, the petalias (an asymmetry between the left and right lobes) suggests a diverge event sometime during the brain expansion after H. habilis.

Mechanisms Responsible for the Difference: 

The mechanisms allowing for greater gyrification in humans is not completely understood, in part owing to the complexities of neuronal migration and brain development. The reelin gene, important for normal neuronal migration, may be involved. Mutations in the reelin gene are associated with lissencephaly, autism, and schizophrenia (Kippenhan et al., 2005; Palaniyappan et al., 2012).

Implications for Understanding Modern Humans: 

Increased gyrification allows for greater overall number of neurons and greater processing efficiency. Understanding the mechanism by which humans have achieved the highest gyrification index may give clues on how humans achieve their remarkable cognitive abilities, as well as provide insights into neuronal developmental disorders, such as the widely prevalent autism spectrum disorders. Further, differences in the patterning of gyrification may signify underlying changes in connectivity between different areas of the brain.

Occurrence in Other Animals: 

Gyrification in general is not unique among humans. The gyrification index increases from prosimians to humans.

References

  1. Growth and folding of the mammalian cerebral cortex: from molecules to malformations., Sun, Tao, and Hevner Robert F. , Nat Rev Neurosci, 04/2014, Volume 15, Issue 4, p.217-32, (2014)
  2. Increased gyrification, but comparable surface area in adolescents with autism spectrum disorders., Wallace, G. L., Robustelli B., Dankner N., Kenworthy L., Giedd J. N., and Martin A. , Brain, 06/2013, Volume 136, Issue Pt 6, p.1956-67, (2013)
  3. Differential effects of surface area, gyrification and cortical thickness on voxel based morphometric deficits in schizophrenia., Palaniyappan, L., and Liddle P. F. , Neuroimage, 03/2012, Volume 60, Issue 1, p.693-9, (2012)
  4. Hominin paleoneurology: where are we now?, Falk, D. , Prog Brain Res, Volume 195, p.255-72, (2012)
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  8. Genetic contributions to human gyrification: sulcal morphometry in Williams syndrome., Kippenhan, J. S., Olsen R. K., Mervis C. B., Morris C. A., Kohn P., Meyer-Lindenberg A., and Berman K. F. , J Neurosci, 08/2005, Volume 25, Issue 34, p.7840-6, (2005)
  9. Humans and great apes share a large frontal cortex., Semendeferi, K, Lu A, Schenker N, and Damasio H , Nat Neurosci, 03/2002, Volume 5, Issue 3, p.272-6, (2002)
  10. The failure of the gyrification index (GI) to account for volumetric reorganization in the evolution of the human brain, Holloway, R. L. , Journal of Human Evolution, 03/1992, Volume 22, Issue 3, p.163 - 170, (1992)
  11. Cortical folding, the lunate sulcus and the evolution of the human brain, Armstrong, E., Zilles K., Curtis M., and Schleicher A. , 04/1991, Volume 20, Issue 4, p.341 - 348, (1991)
  12. Gyrification in the cerebral cortex of primates., Zilles, K, Armstrong E, Moser K H., Schleicher A, and Stephan H , Brain Behav Evol, Volume 34, Issue 3, p.143-50, (1989)
  13. The human pattern of gyrification in the cerebral cortex., Zilles, K, Armstrong E, Schleicher A, and Kretschmann H J. , Anat Embryol (Berl), 1988, Volume 179, Issue 2, p.173-9, (1988)