Brain Structure and Function

, Volume 224, Issue 5, pp 1933–1946 | Cite as

The motor cortex of the sheep: laminar organization, projections and diffusion tensor imaging of the intracranial pyramidal and extrapyramidal tracts

  • Antonella Peruffo
  • Livio Corain
  • Cristiano Bombardi
  • Cinzia Centelleghe
  • Enrico Grisan
  • Jean-Marie Graïc
  • Pietro Bontempi
  • Annamaria Grandis
  • Bruno CozziEmail author
Original Article


The laminar organization of the motor cortex of the sheep and other large domestic herbivores received scarce attention and is generally considered homologous to that of rodents and primates. Thickness of the cortex, subdivision into layers and organization are scarcely known. In the present study, we applied different modern morphological, mathematical and image-analyses techniques to the study of the motor area that controls movements of the forelimb in the sheep. The thickness of the cortex resulted comparable to that of other terrestrial Cetartiodactyls (but thicker than in marine Cetartiodactyls of similar body mass). The laminar organization showed marked development of layer 1, virtual absence of layer 4, and image analysis suggested prevalence of large irregular neural cells in the deeper layers. Diffusion tensor imaging revealed robust projections from the motor cortex to the pyramids in the brainstem, and well evident tracts descending to the tegmentum of the mesencephalon and dorsal pons. Our data contrast the general representation of the motor system of this species, considered to be predominantly based on extra-pyramidal tracts that originate from central pattern generators in the brainstem.


Motor cortex Sheep Laminar organization Cortical projections 



This study was funded by Grant # 2015Y5W9YP from the Italian Ministry of Education, University and Research to BC, including also LC, AP, CB, EG, and J-MG.


This study was funded by Grant # 2015Y5W9YP from the Italian Ministry of Education, University and Research to BC, including also LC, AP, CB, EG, and J-MG.

Compliance with ethical standards

Conflict of interest

The authors declare no conflict of interest.

Statement on welfare of animals

All the brains used in the present study were collected from sheep slaughtered at commercial abattoirs for meat production and commerce. Under these conditions, no ethical approval is required. However, the sheep were treated according to the European Community Council directive (86/609/EEC) concerning animal welfare during the commercial slaughtering process, and constantly monitored under mandatory official veterinary medical care. Furthermore, although not required for this study, the national ethical commission (Ministry of health authorization n_457/2016-PR) approved the general study design of the funded project (Grant # 2015Y5W9YP from the Italian Ministry of Education, University and Research to BC, including also LC, AP, CB, EG, and J-MG.), which was created in compliance with Italian legislation on animal experiments.

Supplementary material

429_2019_1885_MOESM1_ESM.csv (18.4 mb)
Supplementary material 1 (CSV 18802 kb)
429_2019_1885_MOESM2_ESM.docx (16 kb)
Statistical design and data analytics (DOCX 15 kb)
429_2019_1885_MOESM3_ESM.avi (30.5 mb)
3D DTI representation of the corticospinal projections of the motor cortex of the sheep (AVI 31257 kb)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  • Antonella Peruffo
    • 1
  • Livio Corain
    • 2
  • Cristiano Bombardi
    • 3
  • Cinzia Centelleghe
    • 1
  • Enrico Grisan
    • 4
    • 5
  • Jean-Marie Graïc
    • 1
  • Pietro Bontempi
    • 6
  • Annamaria Grandis
    • 3
  • Bruno Cozzi
    • 1
    Email author
  1. 1.Department of Comparative Biomedicine and Food ScienceUniversity of PadovaLegnaroItaly
  2. 2.Department of Management and EngineeringUniversity of PadovaVicenzaItaly
  3. 3.Department of Veterinary Medical SciencesUniversity of BolognaOzzano dell’EmiliaItaly
  4. 4.Department of Information EngineeringUniversity of PadovaPadovaItaly
  5. 5.Department of Biomedical EngineeringKing’s CollegeLondonUK
  6. 6.Department of Computer ScienceUniversity of VeronaVeronaItaly

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