Architectonic subdivisions of neocortex in the tree shrew (Tupaia belangeri)

Anat Rec (Hoboken). 2009 Jul;292(7):994-1027. doi: 10.1002/ar.20916.


Tree shrews are small mammals that bear some semblance to squirrels, but are actually close relatives of primates. Thus, they have been extensively studied as a model for the early stages of primate evolution. In this study, subdivisions of cortex were reconstructed from brain sections cut in the coronal, sagittal, or horizontal planes, and processed for parvalbumin, SMI-32-immunopositive neurofilament protein epitopes, vesicle glutamate transporter 2 (VGluT2), free ionic zinc, myelin, cytochrome oxidase, and Nissl substance. These different procedures revealed similar boundaries between areas, suggesting the detection of functionally relevant borders and allowed a more precise demarcation of cortical areal boundaries. Primary cortical areas were most clearly revealed by the zinc stain, because of the poor staining of layer 4, as thalamocortical terminations lack free ionic zinc. Area 17 (V1) was especially prominent, as the broad layer 4 was nearly free of zinc stain. However, this feature was less pronounced in primary auditory and somatosensory cortex. In primary sensory areas, thalamocortical terminations in layer 4 densely express VGluT2. Auditory cortex consists of two architectonically distinct subdivisions, a primary core region (Ac), surrounded by a belt region (Ab) that had a slightly less developed koniocellular appearance. Primary motor cortex (M1) was identified by the absence of VGluT2 staining in the poorly developed granular layer 4 and the presence of SMI-32-labeled pyramidal cells in layers 3 and 5. The presence of well-differentiated cortical areas in tree shrews indicates their usefulness in studies of cortical organization and function.

Publication types

  • Research Support, N.I.H., Extramural

MeSH terms

  • Animals
  • Auditory Cortex / anatomy & histology
  • Auditory Cortex / physiology
  • Biological Evolution
  • Brain Mapping / methods*
  • Classification
  • Coloring Agents
  • Electron Transport Complex IV / metabolism
  • Immunohistochemistry
  • Motor Cortex / anatomy & histology
  • Motor Cortex / physiology
  • Myelin Proteins / metabolism
  • Neocortex / anatomy & histology*
  • Neocortex / physiology
  • Neurofilament Proteins / metabolism
  • Nissl Bodies / ultrastructure
  • Parvalbumins / metabolism
  • Phylogeny
  • Pyramidal Cells / cytology
  • Pyramidal Cells / physiology
  • Somatosensory Cortex / anatomy & histology
  • Somatosensory Cortex / physiology
  • Species Specificity
  • Tupaia / anatomy & histology*
  • Tupaia / physiology
  • Vesicular Glutamate Transport Protein 2 / metabolism
  • Visual Cortex / anatomy & histology
  • Visual Cortex / physiology
  • Zinc / metabolism


  • Coloring Agents
  • Myelin Proteins
  • Neurofilament Proteins
  • Parvalbumins
  • Vesicular Glutamate Transport Protein 2
  • Electron Transport Complex IV
  • Zinc