TABLE 3.
Studies | Subjects used for Golgi staining | Region and cortical layer in which VENs were analyzed | Golgi staining method | VEN soma morphology | VEN apical dendrite/stem morphology | VEN basal dendrite/stem morphology | VEN axon morphology |
Cajal, 1899 | 1-month-old human female | FI; layer V | Classic Golgi staining | Spindle-like | Prominent with gradual decrease in thickness | Prominent, ends with a dendritic tuft (basilar skirt) | Arises from basal stem/dendrite |
Watson et al., 2006 | 23-year-old human male | ACC and FI; layer V | Modified Golgi technique | Large and elongated, clear demarcation between soma and dendrites is visible in some neurons | Prominent, no additional dendrites or branching for a half-soma’s distance along the length of the proximal dendrites | Prominent, no additional dendrites or branching for a half-soma’s distance along the length of the proximal dendrites, most neurons shown lack a basilar skirt | Not shown |
Evrard et al., 2012 | One rhesus macaque | Anterior insula; layer Vb | Rapid Golgi-Cox | Spindle-like | Branches distally into several thinner spiny dendrites | Branches into thinner spiny dendrites in layer VI, visible basilar skirt | Arises from basal dendrite |
Butti et al., 2014 | One female pygmy hippopotamus | ACC, frontal magnocellular cortex, lateral gyrus*; layer V** | Modified rapid Golgi | Stout, sometimes slender | Almost as thick as the soma | Almost as thick as the soma, occasionally dividing into two branches with a hint of a basilar skirt | Not shown |
Banovac et al., 2019 | 5 adult human male subjects | ACC; layer Vb | Rapid Golgi and Golgi-Cox | Large spindle-like, stick-like or corkscrew-like, on most neurons there is no clear demarcation between soma and dendrites | Very thick origin (up to 8 μm), very gradual decrease in thickness | Constant or very gradual decrease in thickness, low bifurcation degree, but high arborization (brush-like basilar skirt) | Arises from end of basal stem/dendrite |
Correa-Júnior et al., 2020 | 4 adult human subjects (two male and two female) | ACC; layer Vb | “Single-section” Golgi | Large, elongated and spindle-shaped | Prominent with gradual decrease in thickness | Prominent with gradual decrease in thickness, low bifurcation degree, level of arborization depends on subtype (basilar skirt present in some neurons) | Not shown |
*Other cortical areas were also analyzed, but only using Nissl staining or immunohistochemistry (frontopolar, temporal, parietal, and occipital cortex) and VENs were claimed to be found in all the analyzed cortical regions.
**VENs were claimed to occasionally have been found in the deep layer III of numerous cortical areas, but they were still most abundant in layer V and shown on Golgi only in layer V.