Overview	
  of	
  Associational	
  Cortex	
  •	
  Spring	
  2013	
  
1
Medical	
  Neuroscience	
  2013	
  |	
  Tutorial	
  
Overview	
  of	
  Associational	
  Cortex	
  
MAP	
  TO	
  NEUROSCIENCE	
  CORE	
  CONCEPTS
1
	
  
NCC5.	
   Intelligence	
  arises	
  as	
  the	
  brain	
  reasons,	
  plans,	
  and	
  solves	
  problems.	
  
NCC7.	
   The	
  human	
  brain	
  endows	
  us	
  with	
  a	
  natural	
  curiosity	
  to	
  understand	
  how	
  the	
  world	
  works.	
  
LEARNING	
  OBJECTIVES	
  
After	
  study	
  of	
  the	
  assigned	
  learning	
  materials,	
  the	
  student	
  will:	
  
1. Characterize	
  the	
  behaviors	
  and	
  corresponding	
  neural	
  processes	
  that	
  contribute	
  to	
  cognition.	
  
2. Discuss	
  the	
  major	
  similarities	
  and	
  differences	
  in	
  the	
  organization	
  of	
  primary	
  cortex	
  and	
  
associational	
  cortex.	
  
TUTORIAL	
  OUTLINE	
  
I. Introduction	
  to	
  the	
  neuroscience	
  of	
  cognition	
  
A. lay-­‐person’s	
  definition	
  of	
  cognition:	
  the	
  process	
  of	
  knowing	
  
B. neurobiological	
  definition:	
  the	
  neural	
  processes	
  by	
  which	
  the	
  brain	
  integrates	
  
meaningful	
  stimuli,	
  memory,	
  and	
  internal	
  motivations	
  producing	
  perceptional	
  
awareness	
  and	
  appropriate	
  behavior	
  
C. Table	
  1.	
  	
  Neural	
  processes	
  that	
  contribute	
  to	
  cognition	
  
Cognitive	
  
process	
   Metaphor	
  /	
  Example	
   Neural	
  Process	
  
Attention	
   cognitive	
  “search	
  light”	
   modulatory	
  influences	
  of	
  brainstem	
  reticular	
  formation,	
  
hypothalamus	
  and	
  basal	
  forebrain	
  nuclei	
  on	
  thalamic	
  and	
  cortical	
  
processes	
  
Recognition	
   finding	
  a	
  friend’s	
  face	
  in	
  a	
  crowd	
   coding	
  of	
  feature	
  representations	
  in	
  primary	
  and	
  higher	
  order	
  
sensory	
  cortices	
  
Integration	
   knowing	
  that	
  friend	
   integration	
  (“association”)	
  of	
  disparate	
  processing	
  streams	
  in	
  
associational	
  cortices	
  
Planning	
   deciding	
  to	
  seek	
  out	
  that	
  friend	
   processing	
  in	
  executive	
  associational	
  cortices	
  in	
  prefrontal	
  cortex	
  
of	
  frontal	
  lobe	
  
Selection	
  &	
  
execution	
  
Walking	
  towards	
  that	
  friend	
  and	
  
engage	
  in	
  conversation	
  
implementation	
  of	
  short-­‐term	
  and	
  long-­‐term	
  plans	
  via	
  somatic	
  
motor,	
  visceral	
  motor	
  and	
  emotional	
  motor	
  systems	
  
	
  
1
	
   Visit	
  BrainFacts.org	
  for	
  Neuroscience	
  Core	
  Concepts	
  (©2012	
  Society	
  for	
  Neuroscience)	
  that	
  offer	
  fundamental	
  principles	
  
about	
  the	
  brain	
  and	
  nervous	
  system,	
  the	
  most	
  complex	
  living	
  structure	
  known	
  in	
  the	
  universe.	
  
Overview	
  of	
  Associational	
  Cortex	
  •	
  Spring	
  2013	
  
2
II. Anatomy	
  of	
  the	
  Associational	
  Cortex	
  
A. review	
  of	
  cortical	
  structure	
  and	
  function	
  
1. “canonical”	
  (i.e.,	
  standard/representative)	
  cortical	
  microcircuit	
  
a. the	
  columnar	
  circuit	
  is	
  the	
  fundamental	
  unit	
  of	
  processing	
  and	
  
computation	
  in	
  the	
  complex	
  circuitry	
  of	
  the	
  cerebral	
  cortex	
  
b. within	
  a	
  cortical	
  column,	
  different	
  types	
  of	
  excitatory	
  and	
  inhibitory	
  
neurons	
  populate	
  distinct	
  cellular	
  layers	
  
(i) the	
  main	
  excitatory	
  neurons	
  are	
  pyramidal	
  neurons	
  
(ii) the	
  inhibitory	
  neurons	
  are	
  diverse	
  morphologically	
  and	
  
functionally	
  
c. each	
  layer	
  maintains	
  distinct	
  sets	
  of	
  inputs	
  and	
  outputs	
  
(i) pyramidal	
  neurons	
  project:	
  
- to	
  other	
  neurons	
  in	
  the	
  same	
  layer	
  and	
  across	
  layers	
  
within	
  the	
  local	
  column	
  
- to	
  other	
  columns	
  in	
  the	
  same	
  cortical	
  area	
  
- to	
  other	
  columns	
  in	
  other	
  cortical	
  areas,	
  including	
  the	
  
corresponding	
  area	
  in	
  the	
  opposite	
  cerebral	
  hemisphere	
  
- to	
  subcortical	
  targets	
  (e.g.,	
  corticospinal	
  tract	
  neurons)	
  
(ii) most	
  inhibitory	
  neurons	
  project	
  locally	
  within	
  their	
  home	
  layer	
  
and	
  column,	
  but	
  some	
  do	
  project	
  across	
  cortical	
  columns	
  within	
  
the	
  same	
  area	
  (few	
  project	
  to	
  other	
  areas;	
  very	
  few	
  if	
  any	
  
project	
  to	
  the	
  other	
  hemisphere	
  or	
  subcortically)	
  
B. distribution	
  of	
  associational	
  cortex	
  (see	
  Figure	
  26.12
)	
  
1. only	
  about	
  25%	
  of	
  the	
  cerebral	
  mantle	
  is	
  comprised	
  of	
  primary	
  sensory	
  and	
  
motor	
  cortex;	
  the	
  remaining	
  75%	
  is	
  termed	
  associational	
  cortex	
  
2. with	
  the	
  possible	
  exception	
  of	
  the	
  occipital	
  lobe	
  (which	
  is	
  largely	
  if	
  not	
  entirely	
  
devoted	
  to	
  increasingly	
  complex	
  aspects	
  of	
  visual	
  processing),	
  all	
  other	
  lobes	
  
contain	
  associational	
  cortex	
  
3. as	
  its	
  name	
  implies,	
  the	
  associational	
  cortex	
  integrates	
  (or	
  associates)	
  different	
  
types	
  of	
  inputs,	
  some	
  of	
  which	
  come	
  from	
  primary	
  and	
  higher	
  order	
  sensory	
  
cortex	
  
4. it	
  is	
  in	
  the	
  associational	
  cortices	
  that	
  most	
  of	
  the	
  neural	
  processes	
  of	
  cognition	
  
are	
  carried	
  out	
  (together	
  with	
  interactions	
  among	
  subcortical	
  neural	
  centers)	
  
C. differences	
  between	
  associational	
  cortex	
  and	
  primary	
  cortex	
  (see	
  Figure	
  26.4)	
  
1. unlike	
  primary	
  cortices,	
  which	
  receive	
  their	
  main	
  input	
  from	
  specific	
  sensory	
  and	
  
motor	
  relay	
  nuclei	
  of	
  the	
  thalamus,	
  the	
  associational	
  cortex	
  receives	
  its	
  most	
  
influential	
  input	
  from	
  other	
  parts	
  of	
  the	
  cortex	
  
2
	
  Figure	
  references	
  to	
  Purves	
  et	
  al.,	
  Neuroscience,	
  5
th
	
  Ed.,	
  Sinauer	
  Assoc.,	
  Inc.,	
  2012.	
  [click	
  here]	
  
Overview	
  of	
  Associational	
  Cortex	
  •	
  Spring	
  2013	
  
3
2. associational	
  cortex	
  does	
  receive	
  input	
  from	
  thalamus,	
  but	
  it	
  arises	
  from	
  
thalamic	
  nuclei	
  (pulvinar,	
  posterior	
  nuclei,	
  and	
  mediodorsal	
  nucleus)	
  that	
  are	
  
themselves	
  driven	
  by	
  highly	
  processed	
  “feedback”	
  projections	
  from	
  the	
  cortex	
  
(rather	
  than	
  from	
  “feedforward”	
  sensory	
  systems)	
  (see	
  also	
  Appendix	
  BoxA)	
  
3. other	
  sources	
  of	
  input	
  come	
  from	
  brainstem	
  modulatory	
  systems:	
  
a. noradrenergic	
  cells	
  of	
  the	
  locus	
  coeruleus	
  
b. serotonergic	
  cells	
  of	
  the	
  Raphe	
  nuclei	
  
c. dopaminergic	
  cells	
  of	
  the	
  ventral	
  tegmental	
  area)	
  
4. and	
  from	
  the	
  basal	
  forebrain:	
  
a. cholinergic	
  cells	
  in	
  the	
  basal	
  nucleus	
  of	
  Meynert	
  
	
  
STUDY	
  QUESTION	
  
Cognition	
  is	
  an	
  emergent	
  property	
  of	
  brain	
  function,	
  reflecting	
  (as	
  Charles	
  Sherrington	
  famously	
  put	
  it	
  a	
  
century	
  ago)	
  the	
  integrative	
  action	
  of	
  the	
  nervous	
  system.	
  	
  Nevertheless,	
  it	
  is	
  useful	
  to	
  identify	
  and	
  study	
  
distinct	
  neurobiological	
  mechanisms	
  that	
  support	
  disparate	
  functions	
  that	
  contribute	
  to	
  cognition.	
  	
  
Which	
  cognitive	
  process	
  is	
  attributable	
  to	
  the	
  functions	
  of	
  brainstem	
  modulatory	
  (biogenic	
  amine)	
  
systems?	
  
A. selection	
  and	
  execution	
  of	
  short	
  and	
  long-­‐term	
  plans	
  
B. integration	
  of	
  disparate	
  processing	
  streams	
  
C. recognition	
  and	
  coding	
  of	
  salient	
  features	
  
D. attentional	
  modulation	
  of	
  sensory	
  processing	
  
E. planning	
  appropriate	
  behaviors	
  
	
  

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lecture of Associational Cortex in detail.pdf

  • 1. Overview  of  Associational  Cortex  •  Spring  2013   1 Medical  Neuroscience  2013  |  Tutorial   Overview  of  Associational  Cortex   MAP  TO  NEUROSCIENCE  CORE  CONCEPTS 1   NCC5.   Intelligence  arises  as  the  brain  reasons,  plans,  and  solves  problems.   NCC7.   The  human  brain  endows  us  with  a  natural  curiosity  to  understand  how  the  world  works.   LEARNING  OBJECTIVES   After  study  of  the  assigned  learning  materials,  the  student  will:   1. Characterize  the  behaviors  and  corresponding  neural  processes  that  contribute  to  cognition.   2. Discuss  the  major  similarities  and  differences  in  the  organization  of  primary  cortex  and   associational  cortex.   TUTORIAL  OUTLINE   I. Introduction  to  the  neuroscience  of  cognition   A. lay-­‐person’s  definition  of  cognition:  the  process  of  knowing   B. neurobiological  definition:  the  neural  processes  by  which  the  brain  integrates   meaningful  stimuli,  memory,  and  internal  motivations  producing  perceptional   awareness  and  appropriate  behavior   C. Table  1.    Neural  processes  that  contribute  to  cognition   Cognitive   process   Metaphor  /  Example   Neural  Process   Attention   cognitive  “search  light”   modulatory  influences  of  brainstem  reticular  formation,   hypothalamus  and  basal  forebrain  nuclei  on  thalamic  and  cortical   processes   Recognition   finding  a  friend’s  face  in  a  crowd   coding  of  feature  representations  in  primary  and  higher  order   sensory  cortices   Integration   knowing  that  friend   integration  (“association”)  of  disparate  processing  streams  in   associational  cortices   Planning   deciding  to  seek  out  that  friend   processing  in  executive  associational  cortices  in  prefrontal  cortex   of  frontal  lobe   Selection  &   execution   Walking  towards  that  friend  and   engage  in  conversation   implementation  of  short-­‐term  and  long-­‐term  plans  via  somatic   motor,  visceral  motor  and  emotional  motor  systems     1   Visit  BrainFacts.org  for  Neuroscience  Core  Concepts  (©2012  Society  for  Neuroscience)  that  offer  fundamental  principles   about  the  brain  and  nervous  system,  the  most  complex  living  structure  known  in  the  universe.  
  • 2. Overview  of  Associational  Cortex  •  Spring  2013   2 II. Anatomy  of  the  Associational  Cortex   A. review  of  cortical  structure  and  function   1. “canonical”  (i.e.,  standard/representative)  cortical  microcircuit   a. the  columnar  circuit  is  the  fundamental  unit  of  processing  and   computation  in  the  complex  circuitry  of  the  cerebral  cortex   b. within  a  cortical  column,  different  types  of  excitatory  and  inhibitory   neurons  populate  distinct  cellular  layers   (i) the  main  excitatory  neurons  are  pyramidal  neurons   (ii) the  inhibitory  neurons  are  diverse  morphologically  and   functionally   c. each  layer  maintains  distinct  sets  of  inputs  and  outputs   (i) pyramidal  neurons  project:   - to  other  neurons  in  the  same  layer  and  across  layers   within  the  local  column   - to  other  columns  in  the  same  cortical  area   - to  other  columns  in  other  cortical  areas,  including  the   corresponding  area  in  the  opposite  cerebral  hemisphere   - to  subcortical  targets  (e.g.,  corticospinal  tract  neurons)   (ii) most  inhibitory  neurons  project  locally  within  their  home  layer   and  column,  but  some  do  project  across  cortical  columns  within   the  same  area  (few  project  to  other  areas;  very  few  if  any   project  to  the  other  hemisphere  or  subcortically)   B. distribution  of  associational  cortex  (see  Figure  26.12 )   1. only  about  25%  of  the  cerebral  mantle  is  comprised  of  primary  sensory  and   motor  cortex;  the  remaining  75%  is  termed  associational  cortex   2. with  the  possible  exception  of  the  occipital  lobe  (which  is  largely  if  not  entirely   devoted  to  increasingly  complex  aspects  of  visual  processing),  all  other  lobes   contain  associational  cortex   3. as  its  name  implies,  the  associational  cortex  integrates  (or  associates)  different   types  of  inputs,  some  of  which  come  from  primary  and  higher  order  sensory   cortex   4. it  is  in  the  associational  cortices  that  most  of  the  neural  processes  of  cognition   are  carried  out  (together  with  interactions  among  subcortical  neural  centers)   C. differences  between  associational  cortex  and  primary  cortex  (see  Figure  26.4)   1. unlike  primary  cortices,  which  receive  their  main  input  from  specific  sensory  and   motor  relay  nuclei  of  the  thalamus,  the  associational  cortex  receives  its  most   influential  input  from  other  parts  of  the  cortex   2  Figure  references  to  Purves  et  al.,  Neuroscience,  5 th  Ed.,  Sinauer  Assoc.,  Inc.,  2012.  [click  here]  
  • 3. Overview  of  Associational  Cortex  •  Spring  2013   3 2. associational  cortex  does  receive  input  from  thalamus,  but  it  arises  from   thalamic  nuclei  (pulvinar,  posterior  nuclei,  and  mediodorsal  nucleus)  that  are   themselves  driven  by  highly  processed  “feedback”  projections  from  the  cortex   (rather  than  from  “feedforward”  sensory  systems)  (see  also  Appendix  BoxA)   3. other  sources  of  input  come  from  brainstem  modulatory  systems:   a. noradrenergic  cells  of  the  locus  coeruleus   b. serotonergic  cells  of  the  Raphe  nuclei   c. dopaminergic  cells  of  the  ventral  tegmental  area)   4. and  from  the  basal  forebrain:   a. cholinergic  cells  in  the  basal  nucleus  of  Meynert     STUDY  QUESTION   Cognition  is  an  emergent  property  of  brain  function,  reflecting  (as  Charles  Sherrington  famously  put  it  a   century  ago)  the  integrative  action  of  the  nervous  system.    Nevertheless,  it  is  useful  to  identify  and  study   distinct  neurobiological  mechanisms  that  support  disparate  functions  that  contribute  to  cognition.     Which  cognitive  process  is  attributable  to  the  functions  of  brainstem  modulatory  (biogenic  amine)   systems?   A. selection  and  execution  of  short  and  long-­‐term  plans   B. integration  of  disparate  processing  streams   C. recognition  and  coding  of  salient  features   D. attentional  modulation  of  sensory  processing   E. planning  appropriate  behaviors