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Insula
The Hidden Cortex


       Rebecca E. Martin
         January, 2009
Insula
The Hidden Cortex
The insula is the cortical part of the brain
that is not visible when looking at the
surface of the brain.


                 Rebecca E. Martin
                   January, 2009
A?
       UL
    INS
the
A?
       UL
    INS
the
 Anatomy
Insula is Latin for
Insula is Latin for
    ISLAND
The Insula Looks Like a
                Seashell                                                  FIG 2. The transverse gyrus and the orbitofrontal insula rela-
                                                                          tionship. Base view of the orbital surface of the left frontal lobe
                                                                          after resection of part of the gyrus rectus (GR) and the anterior
                                                                          portion of the temporal lobe (T). Medial is to the reader’s left. I
                                                                          indicates olfactory bulb; II, optic chiasm and tract. The H-shaped
                                                                          orbital sulcus (arrowheads) defines the MOG, POG, anterior
                                                                          orbital gyrus (AOG), and lateral orbital gyrus (LOG). At the pos-
                                                                          teromedial aspect of the orbitofrontal surface, the posterior por-
                                                                          tion of the MOG merges with the medial portion of the POG to
                                                                          form the prominent PMOL. PMOL gives rise to the transverse
                                                                          insular gyrus (t) that extends laterally to form the pole (po) of the
                                                                          insula just anteroinferomedial to the apex (asterisk) of the insula.
                                                                          The CS (dashed line) curves inferiorly immediately behind and
                                                                          below the apex and the pole en route to join the stem of the
                                                                          sylvian fissure. The ASG (a) and the PSG (p) converge to form the
                                                                          apex of the insula anterior to the CS. In this image, the deliberate
                                                                          slight rotation used to illustrate the course of the transverse
                                                                          gyrus from the PMOL to the pole also rotates the apex medially,
                                                                          so the apex does not appear to lie as far lateral in position as it
                                                                          would in a true base view.


                                                                          cases, including two in which the MSG was equal to
                                                                          the ASG in size).
FIG 1. A–C, Anatomy of the convexity surface of the left insular
                                                                             The PSG formed the precentral gyrus of the insula
lobe after resection of the overlying opercula, vessels, and pia-
                                                                          immediately anterior to the CS in all cases. Typically,
arachnoid. Gross anatomic specimens from a 50-year-old woman
(A), a 63-year-old man (B), and a 71-year-old man (C). The anterior       the short insular sulcus separated the ASG from the
PS (APS), superior PS (SPS), and inferior PS (IPS) define the base
                                                                          MSG, whereas the precentral sulcus of the insula
of the insular lobe. The oblique CS of the insula subdivides the
                                                                          separated the MSG from the PSG (Table 2). In one
insula into larger anterior and smaller posterior lobules.
                                                                          case with absent MSG, the short insular sulcus and
   Anterior lobule: The ASG (a), MSG (m), and PSG (p) form most
of the convexity surface of the anterior lobule. The short insular        the precentral sulcus merged completely, so they
sulcus (SIS) separates ASG from MSG. The precentral sulcus
                                                                          could not be identified as separate structures.
(preCS) separates MSG from PSG. MSG shows variable size and
                                                                             The apex of the insula was formed by the short gyri
depression below the surface of the insula. The number of gyri
                                                                                                                               Naidich et al., 2004
                                                                          in all cases, but the specific pattern of the short gyri
on the convexity surface of the anterior lobule varies substan-
tially: two in A, three in B, and four in C. The apex (asterisk) of the   and the contribution of each short gyrus to the apex
insula forms by the convergence of the inferior ends of some or
                                                                          were variable (Fig 1). The ASG contributed to the
Central   Limen
Insular
Sulcus
                  Gray’s Anatomy
Anterior
connects to
   limbic
structures
     i.e.
    amyg
 thalamus
hippocamp




              Central   Limen
              Insular
              Sulcus
                                Gray’s Anatomy
Anterior
connects to
   limbic
structures
     i.e.
    amyg
 thalamus
hippocamp




                                       Posterior
                                   associated with
              Central   Limen    somatosensory areas,
              Insular              pain pathways
              Sulcus
                                Gray’s Anatomy
The Insula’s
Neighborhood
It is covered by the operculum
It is covered by the operculum
It is covered by the operculum
It is covered by the operculum
It is covered by the operculum
             Thanks for the pics, Dianne and Hendrik!
Adjacent to the lateral ventricle
Insula




         REVIEW!   TÜRE et al., 1999
Insula




         REVIEW!   TÜRE et al., 1999
Insula




         REVIEW!   TÜRE et al., 1999
Insula




         REVIEW!   TÜRE et al., 1999
Insula




         REVIEW!   TÜRE et al., 1999
Insula




         REVIEW!   TÜRE et al., 1999
Insula




         REVIEW!   TÜRE et al., 1999
Medial to the Insula   TÜRE et al., 2000
Vascularization
              TÜRE et al., 2000
Vascularization
              TÜRE et al., 2000
Vascularization
              TÜRE et al., 2000
Vascularization
The Insula is fed entirely by the middle cerebral artery   TÜRE et al., 2000
More Arteries
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Embryology   Afif et al., 2007
Architecture and Histology
                  Chikama, et al., 19970
Anterior
 agranular




                                          Posterior
                                          granular
             Central   Limen
             Insular
                       dysgranular
             Sulcus
                                     Gray’s Anatomy
granular



                               dysgranular



                               agranular


granular - 6 layers
dysgranular - 5-ish layers
agranular - 5 or less layers
                                             Augustine, 1996
granular



                               dysgranular



                               agranular
                                            Wow, that’s similar
                                           to the OFC and the
granular - 6 layers                             Cingulate!
dysgranular - 5-ish layers
agranular - 5 or less layers
                                             Augustine, 1996
Spindle Cells




Phylogeny       Manger, et al., 1998
Spindle Cells
            are only found in
            cingulate and insular
            regions


Phylogeny         Manger, et al., 1998
Connectivity
Major Fiber Tracts Surrounding Insula




                                    TÜRE et al., 1999
Major Fiber Tracts Surrounding Insula




                                    TÜRE et al., 1999
Major Fiber Tracts Surrounding Insula




                                    TÜRE et al., 1999
EFFERENT CONNECTIONS
              Augustine, 1996
AFFERENT CONNECTIONS
              Augustine, 1996
Augustine, 1996
Augustine, 1996
Function
                              gustatory
  vomiting!

                                  cravings


                                     singing!
                                     auditory

                               visual - awareness,
                                   monitoring
subjective feeling
                            autonomic (e.g.
                            cardiac rhythm)

                             NYTimes, 2007
But Also...
Olfactory (close to
olfactory bulbs!)

Post-stroke recovery of
motor function

Associated with neglect

Thermal heat pain
Connectivity Between the Amygdala and Insula




                                     Morris, 2002
A Primary Gustatory Area



Taste (lesion studies
indicate taste intensity
and recognition problems)
A Primary Gustatory Area



Taste (lesion studies
indicate taste intensity
and recognition problems)

Fun fact: Taste info from
both sides of mouth
passes though left insula
Activation Studies

The Insula is:
A Visceral Sensory Area
 Role in integration of
 multimodal sensory
 information, gut feelings,
 disgust, temperature
 perception, pain recognition

 Interoception i.e. “How We
 Feel”
Interoception




◦ subjective feelings, emotion, self-awareness (Craig, 2009, 2002;
  Critchley, 2004)
View of Insula and Uncus




                           TÜRE et al., 1999
The (left) Insula is also:
Involved in Language and Memory

  Part of an “articulatory loop”

    Coordinates speech articulation

  Singing!

    (Right insula may mediate control of vocal tract)

  Insula lesions have been noted to cause aphasia

  Part of network involved in verbal memory - mental navigation
  along memorized routes (hippocampus -> precuneus -> insula)

  Humor (particularly gag and language humor, Watson et al., 2006)
Lesion Studies

                 The Insula:
Is associated with certain types of
mental illness
  Early Alzheimers (atrophy)

  Schizophrenia (abnormalities, e.g. decreased
  grey matter)

  OCD, PTSD, Anxiety, Depression

May Play a Role in Addiction
  Conscious urges to take drugs (Naqvi,
  Bechara, Damasio, 2008)
Proposed Drug Craving Pathways
References
Afif, A., Bouvier, R., Buenerd, A., Trouillas, J., & Mertens, P. (2007). Development of the human fetal insular cortex:
      study of the gyration from 13 to 28 gestational weeks. Brain Structure and Function, 212(3), 335-346.
Augustine, J. R. (1996). Circuitry and functional aspects of the insular lobe in primates
      including humans. Brain Research Reviews, 22(3), 229-244.
Augustine, J. R. (1985). The insular lobe in primates including humans. Neurological Research, 7(1), 2-10.
Balleine, B. W., & Dickinson, A. (2000). The Effect of Lesions of the Insular Cortex on Instrumental Conditioning:
      Evidence for a Role in Incentive Memory. J. Neurosci., 20(23).
Cereda, C., Ghika, J., Maeder, P., & Bogousslavsky, J. (2002). Strokes restricted to the insular cortex. Neurology,
      59(12), 1950-1955.
Craig, A. (. (2003). Interoception: the sense of the physiological condition of the body. Current Opinion in
      Neurobiology, 13(4), 500-505.
Critchley, H. D., Wiens, S., Rotshtein, P., Ohman, A., & Dolan, R. J. (2004). Neural systems supporting interoceptive
      awareness. Nat Neurosci, 7(2), 189-195.
Manger, P., Sum, M., Szymanski, M., Ridgway, S., & Krubitzer, L. (1998). Modular Subdivisions of Dolphin Insular
      Cortex: Does Evolutionary History Repeat Itself? Journal of Cognitive Neuroscience, 10(2), 153-166.
Morris, J. (2002). How do you feel? Trends in Cognitive Sciences, 6(8), 317-319.
Nagai, M., Kishi, K., & Kato, S. (2007). Insular cortex and neuropsychiatric disorders: A review of recent literature.
      European Psychiatry, 22(6), 387-394.
Naidich, T. P., Kang, E., Fatterpekar, G. M., Delman, B. N., Gultekin, S. H., Wolfe, D., et al. (2004). The Insula:
      Anatomic Study and MR Imaging Display at 1.5 T. AJNR Am J Neuroradiol, 25(2), 222-232.
Naqvi, N. H., Rudrauf, D., Damasio, H., & Bechara, A. (2007). Damage to the Insula Disrupts Addiction to Cigarette
      Smoking. Science, 315(5811), 531-534.
Oppenheimer, S. (2006). Cerebrogenic cardiac arrhythmias: Clinical Autonomic Research. Clin Auton Res, 16(1),
     6-11.
Türe U, Yasargil DCH, Al-Mefty O, Yasargil MG. (1999). Topographic anatomy of the insular region. J Neurosurg,
     90, 720 –733
Türe U, Yasargil MG, Al-Mefty O, Yasargil DCH. (2000). Arteries of the insula. J Neurosurg, 92:676 – 687
Thanks!




          NYTimes, 2007

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The Hidden Cortex: Insula Functions and Anatomy

  • 1. Insula The Hidden Cortex Rebecca E. Martin January, 2009
  • 2. Insula The Hidden Cortex The insula is the cortical part of the brain that is not visible when looking at the surface of the brain. Rebecca E. Martin January, 2009
  • 3. A? UL INS the
  • 4. A? UL INS the Anatomy
  • 6. Insula is Latin for ISLAND
  • 7. The Insula Looks Like a Seashell FIG 2. The transverse gyrus and the orbitofrontal insula rela- tionship. Base view of the orbital surface of the left frontal lobe after resection of part of the gyrus rectus (GR) and the anterior portion of the temporal lobe (T). Medial is to the reader’s left. I indicates olfactory bulb; II, optic chiasm and tract. The H-shaped orbital sulcus (arrowheads) defines the MOG, POG, anterior orbital gyrus (AOG), and lateral orbital gyrus (LOG). At the pos- teromedial aspect of the orbitofrontal surface, the posterior por- tion of the MOG merges with the medial portion of the POG to form the prominent PMOL. PMOL gives rise to the transverse insular gyrus (t) that extends laterally to form the pole (po) of the insula just anteroinferomedial to the apex (asterisk) of the insula. The CS (dashed line) curves inferiorly immediately behind and below the apex and the pole en route to join the stem of the sylvian fissure. The ASG (a) and the PSG (p) converge to form the apex of the insula anterior to the CS. In this image, the deliberate slight rotation used to illustrate the course of the transverse gyrus from the PMOL to the pole also rotates the apex medially, so the apex does not appear to lie as far lateral in position as it would in a true base view. cases, including two in which the MSG was equal to the ASG in size). FIG 1. A–C, Anatomy of the convexity surface of the left insular The PSG formed the precentral gyrus of the insula lobe after resection of the overlying opercula, vessels, and pia- immediately anterior to the CS in all cases. Typically, arachnoid. Gross anatomic specimens from a 50-year-old woman (A), a 63-year-old man (B), and a 71-year-old man (C). The anterior the short insular sulcus separated the ASG from the PS (APS), superior PS (SPS), and inferior PS (IPS) define the base MSG, whereas the precentral sulcus of the insula of the insular lobe. The oblique CS of the insula subdivides the separated the MSG from the PSG (Table 2). In one insula into larger anterior and smaller posterior lobules. case with absent MSG, the short insular sulcus and Anterior lobule: The ASG (a), MSG (m), and PSG (p) form most of the convexity surface of the anterior lobule. The short insular the precentral sulcus merged completely, so they sulcus (SIS) separates ASG from MSG. The precentral sulcus could not be identified as separate structures. (preCS) separates MSG from PSG. MSG shows variable size and The apex of the insula was formed by the short gyri depression below the surface of the insula. The number of gyri Naidich et al., 2004 in all cases, but the specific pattern of the short gyri on the convexity surface of the anterior lobule varies substan- tially: two in A, three in B, and four in C. The apex (asterisk) of the and the contribution of each short gyrus to the apex insula forms by the convergence of the inferior ends of some or were variable (Fig 1). The ASG contributed to the
  • 8. Central Limen Insular Sulcus Gray’s Anatomy
  • 9. Anterior connects to limbic structures i.e. amyg thalamus hippocamp Central Limen Insular Sulcus Gray’s Anatomy
  • 10. Anterior connects to limbic structures i.e. amyg thalamus hippocamp Posterior associated with Central Limen somatosensory areas, Insular pain pathways Sulcus Gray’s Anatomy
  • 12. It is covered by the operculum
  • 13. It is covered by the operculum
  • 14. It is covered by the operculum
  • 15. It is covered by the operculum
  • 16. It is covered by the operculum Thanks for the pics, Dianne and Hendrik!
  • 17. Adjacent to the lateral ventricle
  • 18. Insula REVIEW! TÜRE et al., 1999
  • 19. Insula REVIEW! TÜRE et al., 1999
  • 20. Insula REVIEW! TÜRE et al., 1999
  • 21. Insula REVIEW! TÜRE et al., 1999
  • 22. Insula REVIEW! TÜRE et al., 1999
  • 23. Insula REVIEW! TÜRE et al., 1999
  • 24. Insula REVIEW! TÜRE et al., 1999
  • 25. Medial to the Insula TÜRE et al., 2000
  • 26. Vascularization TÜRE et al., 2000
  • 27. Vascularization TÜRE et al., 2000
  • 28. Vascularization TÜRE et al., 2000
  • 29. Vascularization The Insula is fed entirely by the middle cerebral artery TÜRE et al., 2000
  • 31. Embryology Afif et al., 2007
  • 32. Embryology Afif et al., 2007
  • 33. Embryology Afif et al., 2007
  • 34. Embryology Afif et al., 2007
  • 35. Embryology Afif et al., 2007
  • 36. Embryology Afif et al., 2007
  • 37. Embryology Afif et al., 2007
  • 38. Embryology Afif et al., 2007
  • 39. Embryology Afif et al., 2007
  • 40. Embryology Afif et al., 2007
  • 41. Architecture and Histology Chikama, et al., 19970
  • 42. Anterior agranular Posterior granular Central Limen Insular dysgranular Sulcus Gray’s Anatomy
  • 43. granular dysgranular agranular granular - 6 layers dysgranular - 5-ish layers agranular - 5 or less layers Augustine, 1996
  • 44. granular dysgranular agranular Wow, that’s similar to the OFC and the granular - 6 layers Cingulate! dysgranular - 5-ish layers agranular - 5 or less layers Augustine, 1996
  • 45. Spindle Cells Phylogeny Manger, et al., 1998
  • 46. Spindle Cells are only found in cingulate and insular regions Phylogeny Manger, et al., 1998
  • 48. Major Fiber Tracts Surrounding Insula TÜRE et al., 1999
  • 49. Major Fiber Tracts Surrounding Insula TÜRE et al., 1999
  • 50. Major Fiber Tracts Surrounding Insula TÜRE et al., 1999
  • 51. EFFERENT CONNECTIONS Augustine, 1996
  • 52. AFFERENT CONNECTIONS Augustine, 1996
  • 55. Function gustatory vomiting! cravings singing! auditory visual - awareness, monitoring subjective feeling autonomic (e.g. cardiac rhythm) NYTimes, 2007
  • 56. But Also... Olfactory (close to olfactory bulbs!) Post-stroke recovery of motor function Associated with neglect Thermal heat pain
  • 57. Connectivity Between the Amygdala and Insula Morris, 2002
  • 58. A Primary Gustatory Area Taste (lesion studies indicate taste intensity and recognition problems)
  • 59. A Primary Gustatory Area Taste (lesion studies indicate taste intensity and recognition problems) Fun fact: Taste info from both sides of mouth passes though left insula
  • 60. Activation Studies The Insula is: A Visceral Sensory Area Role in integration of multimodal sensory information, gut feelings, disgust, temperature perception, pain recognition Interoception i.e. “How We Feel”
  • 61. Interoception ◦ subjective feelings, emotion, self-awareness (Craig, 2009, 2002; Critchley, 2004)
  • 62. View of Insula and Uncus TÜRE et al., 1999
  • 63. The (left) Insula is also: Involved in Language and Memory Part of an “articulatory loop” Coordinates speech articulation Singing! (Right insula may mediate control of vocal tract) Insula lesions have been noted to cause aphasia Part of network involved in verbal memory - mental navigation along memorized routes (hippocampus -> precuneus -> insula) Humor (particularly gag and language humor, Watson et al., 2006)
  • 64. Lesion Studies The Insula: Is associated with certain types of mental illness Early Alzheimers (atrophy) Schizophrenia (abnormalities, e.g. decreased grey matter) OCD, PTSD, Anxiety, Depression May Play a Role in Addiction Conscious urges to take drugs (Naqvi, Bechara, Damasio, 2008)
  • 66. References Afif, A., Bouvier, R., Buenerd, A., Trouillas, J., & Mertens, P. (2007). Development of the human fetal insular cortex: study of the gyration from 13 to 28 gestational weeks. Brain Structure and Function, 212(3), 335-346. Augustine, J. R. (1996). Circuitry and functional aspects of the insular lobe in primates including humans. Brain Research Reviews, 22(3), 229-244. Augustine, J. R. (1985). The insular lobe in primates including humans. Neurological Research, 7(1), 2-10. Balleine, B. W., & Dickinson, A. (2000). The Effect of Lesions of the Insular Cortex on Instrumental Conditioning: Evidence for a Role in Incentive Memory. J. Neurosci., 20(23). Cereda, C., Ghika, J., Maeder, P., & Bogousslavsky, J. (2002). Strokes restricted to the insular cortex. Neurology, 59(12), 1950-1955. Craig, A. (. (2003). Interoception: the sense of the physiological condition of the body. Current Opinion in Neurobiology, 13(4), 500-505. Critchley, H. D., Wiens, S., Rotshtein, P., Ohman, A., & Dolan, R. J. (2004). Neural systems supporting interoceptive awareness. Nat Neurosci, 7(2), 189-195. Manger, P., Sum, M., Szymanski, M., Ridgway, S., & Krubitzer, L. (1998). Modular Subdivisions of Dolphin Insular Cortex: Does Evolutionary History Repeat Itself? Journal of Cognitive Neuroscience, 10(2), 153-166. Morris, J. (2002). How do you feel? Trends in Cognitive Sciences, 6(8), 317-319. Nagai, M., Kishi, K., & Kato, S. (2007). Insular cortex and neuropsychiatric disorders: A review of recent literature. European Psychiatry, 22(6), 387-394. Naidich, T. P., Kang, E., Fatterpekar, G. M., Delman, B. N., Gultekin, S. H., Wolfe, D., et al. (2004). The Insula: Anatomic Study and MR Imaging Display at 1.5 T. AJNR Am J Neuroradiol, 25(2), 222-232. Naqvi, N. H., Rudrauf, D., Damasio, H., & Bechara, A. (2007). Damage to the Insula Disrupts Addiction to Cigarette Smoking. Science, 315(5811), 531-534. Oppenheimer, S. (2006). Cerebrogenic cardiac arrhythmias: Clinical Autonomic Research. Clin Auton Res, 16(1), 6-11. Türe U, Yasargil DCH, Al-Mefty O, Yasargil MG. (1999). Topographic anatomy of the insular region. J Neurosurg, 90, 720 –733 Türe U, Yasargil MG, Al-Mefty O, Yasargil DCH. (2000). Arteries of the insula. J Neurosurg, 92:676 – 687
  • 67. Thanks! NYTimes, 2007

Editor's Notes

  1. Talk about anatomy here
  2. •The insular lobe corresponds to the fifth lobe of the brain including Brodmann areas 13 through 16
  3. - The insula is located deep inside the brain, at the base of the Sylvian Fissure. 
  4. •it is the only cortical part of the brain that is not visible when looking at the surface of the brain
  5. •The central insular sulcus, divides the insula into two zones that are unequal in size: the anterior insula (larger) and posterior insula (smaller).
  6. •Anterior= 3-5 short gyri number of gyri are highly varied by person and side of the brain
  7. Posterior = 2 long gyri
  8. •The insula encapsulates the claustrum and putamen (i.e. the lenticular nucleus).
  9. •posterior to the pars triangularis
  10. •Insula adjacent to lateral ventricle except for base of insula (in the C-shaped curve of the ventricle).
  11. •It is separated from the operculum by the anterior, superior, and inferior periinsular sulci.
  12. •It is covered from opercula by a layer of arteries
  13. showing a coronal section of the brain through the foramen of Monro and the amygdala, anterior view.
  14. \"the insula connects with portions of the frontal, parietal, and temporal lobes and with the cingulate gyrus.These investigators have also shown that the basal nuclei (in particular, the tail of the caudate nucleus, the putamen,
  15. and the claustrum), the amygdaloid body and other limbic structures, and the dorsal thalamus also connect with the insula.\"
  16. •Insula adjacent to lateral ventricle except for base of insula (in the C-shaped curve of the ventricle).
  17. The insula and basal gangion areas
  18. axial section of the right cerebral hemisphere through the insula, superior view. Abbreviations with white letters denote sulci and fissures.
  19. -ahg = anterior Heschl’s gyrus;
  20. -bf = body of fornix;
  21. -c = claustrum;
  22. -cg = cingulate gyrus;
  23. -cn = caudate nucleus;
  24. -crf = crus of fornix;
  25. -ec = external capsule;
  26. -exc = extreme capsule;
  27. -gcc = genu of corpus callosum;
  28. -gp = globus pallidus;
  29. -ic = internal capsule;
  30. -p = putamen;
  31. -scc = splenium of corpus callosum;
  32. -sf = sylvian fissure;
  33. -t = thalamus.
  34. Fiber dissection of this area reveals vascularization of the lentiform nuclei (which have been removed) and vascularization of the internal capsule by the LLAs (arrow), which arise from M1 segment. (INTERNAL CAPSULE)
  35. -ac = anterior commissure;
  36. -cis = central insular sulcus;
  37. - ia = insular apex;
  38. -ic = internal capsule;
  39. The insula encapsulates the claustrum and putamen (i.e. the lenticular nucleus).
  40. Insula fed entirely by MCA
  41. 1. The same specimen is shown following removal of the entire opercula with preservation of all segments of the MCA.
  42. 2. The same specimen. The M 3 , M 4 and M 5 segments of the MCA have been removed, and the M 2 segment, which supplies the insula, has been preserved over the insular cortex.
  43. 3. The insula is shown following removal of the frontal, parietal, occipital, and temporal lobes from the periinsular sulci. The arteries of the insula originate from the M2segment. The insulo-opercular arteries (arrows)supply the insula and operculum.
  44. dense wall of arteries running in the lateral fissure. The deep anatomical location of the insular cortex together with the dense sylvian vascularization explains why the insula has rarely been investigated in humans.
  45. Drawings showing that approximately 85 to 90% of insular arteries were short and supplied the insular cortex (i) and extreme capsule; 10% were medium sized and supplied, in addition, the claustrum and external capsule; and the remainder 3 to 5%
  46. were long and extended as far as the corona radiata (cr). a = amygdala; gp = globus pallidus; p = putamen. See previous figure legends for additional abbreviations.
  47. •(Afif et al., 2007) Insula part of the telencephalon.
  48. •surface expansion is more limited than that of the other 4 cortices.
  49. •It is suggested that cortical maturation (sulcation and gyration) and vascularization of the lateral surface of the brain starts with the insular region, suggesting that this region is a central area of cortical development
  50. •Right side develops earlier but left side usually ends up larger in adults (language function?)
  51. •First cortex to grow.  It is anchored to underlying structures so maintains its position while the temporal lobe pivots around it.  Then becomes covered as lobes enlarge.