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Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory, Seoul National University http://bi.snu.ac.kr/

Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

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Prolog : LGN & Ocular Dominance Column at Visual Cortex 3(C) 2009, SNU Biointelligence Lab, Visual Pathway to Primary Visual Cortex for Mammals (http://scienceblogs.com/purepedantry/ 2007/10/ocular_dominance_columns_a nd_t.php) Connection between Retina and LGN (http://dels.nas.edu/ilar_n/ilarjour nal/46_4/html/v4604Kaas.shtml) Retinotopic maps in V1 (http://www.journalofvision.org/3/10/1/article.aspx) Inputs to LGN (Bear et al. Neuroscience: Exploring the brain. (Lippincott Williams & Wilkins: 2006).)

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Page 1: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Ch 5. The Patterning of Neural Connec-tions

5.5 ~ 5.6

Adaptive Cooperative Systems, Martin Beckerman, 1997.

Summarized by Kwonill, Kim

Biointelligence Laboratory, Seoul National Universityhttp://bi.snu.ac.kr/

Page 2: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

2(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Contents

Prolog : LGN & Ocular Dominance Column at Visual Cortex 5.5 The Multiple Constraint Model

¨ 5.5.1 Position-Independent Affinity¨ 5.5.2 Fiber-Fiber Repulsion¨ 5.5.3 Nearest-Neighbor Correlated Activity¨ 5.5.4 Position-Dependent Affinity¨ 5.5.5 Multiple Stable States in the Retinotectal Projection

5.6 Morphogenesis of the Lateral Geniculate Nucleus¨ 5.6.1 Interaction Potentials¨ 5.6.2 Induction of the Laminar Transition¨ 5.6.3 Trapping of the Transition by the Blind Spot

Page 3: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Prolog : LGN & Ocular Dominance Column at Visual Cortex

3(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Visual Pathway to Primary Visual Cor-tex for Mammals(http://scienceblogs.com/purepedantry/2007/10/ocular_dominance_columns_and_t.php)

Connection between Retina and LGN (http://dels.nas.edu/ilar_n/ilarjournal/46_4/html/v4604Kaas.shtml)

Retinotopic maps in V1(http://www.journalofvision.org/3/10/1/article.aspx)

Inputs to LGN(Bear et al. Neuroscience: Exploring the brain. (Lippincott Williams & Wilkins: 2006).)

Page 4: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Prolog : LGN & Ocular Dominance Col-umn at Visual Cortex

4(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Inducing Ocular Dominance Columns by the Transplantation of a third eye (http://www.nature.com/nrn/journal/v3/n1/box/nrn703_BX1.html)

Ocular Dominance Column (https://bbs.stardestroyer.net/view-topic.php?f=5&t=124049&start=0)

Page 5: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Question

What can we infer from these models?

5(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 6: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Multiple Constraint Model

Steinberg’s differential adhesion hypothesis¨ Minimize the adhesive-free energy

Self sorting of cells Morphology or Hierarchy

Multiple constraint model of Fraser & Perkel¨ Influences of

Chemotropic factor Electrical activity Cell surface molecules

¨ Adhesive-free energy Fiber-fiber & fiber-tectum interaction

6(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

E

Page 7: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Arbor Discs & Tectal Disc

Arbor terminal and tectum are modeled as discs¨ Diameter of a arbor disc =

10% of diameter of a tectal disc

7(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Tectum

Arbor Terminal

Arbor Terminal

Arbor Terminal

Page 8: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Position-Independent Affinity

The contribution to the total free energy of the position-independent affinity of the ith fiber

¨ c0: coupling strength (positive)¨ : fractional overlap of the ith fiber disc with the optic tectum

¨ Ignore boundary effects Square potential well

8(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

0 0i iE c (5.1)

i

Page 9: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Fiber-Fiber Repulsion

Short-range fiber-fiber repulsion due to interactions of the ith fiber with the others

¨ c1: positive coupling constant (< c0 )¨ : percent overlap between fibers i and j¨ : the distance between the retinal ganglion

cells responsible for fibers i and j¨ , : constants

9(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

(5.2)

ijr

1a 2a

Page 10: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Nearest-Neighbor Correlated Activity

Electrical-activity-dependent interaction¨ Neighboring retinal ganglion cells

Neighboring tectal cells¨ Weak

10(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 11: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Position-Dependent Affinity

A set of position dependent affinities between growth cones and their tectal targets

Fiber-fiber term

Fiber-tectum term

11(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

2 2 3AP DV

i ij ij ij ijj j

E c r c r (5.3)

2 2 2( ) ( )DV APij ij ijr r r (5.4)

3 4 5(1 ) (1 )AP DVi i i iE c t c t (5.5)

Page 12: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Total Free Energy

¨ Position-Independent Affinity¨ Fiber-Fiber Repulsion

+ Nearest-Neighbor Correlated Activity¨ Fiber-fiber Position-Dependent Affinity¨ Fiber-tectum Position-Dependent Affinity

12(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

0 1 2 3( )tot i i i ii

E E E E E (5.6)

Page 13: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Multiple Stable States in the Retinotectal Projection Simulated Annealing Without the retinal input & the tectal

environment modification¨ Fully satisfying energy constraints¨ Normal projection

With tectal environment modifica-tion¨ Partially satisfying energy constraints¨ Topography (uniform)

13(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 14: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Multiple Stable States in the Retinotectal Projection Ablation

With the retinal inputs¨ Ocular Dominance

14(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 15: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Morphogenesis of the LGN

Visual field of the retina of the rhesus monkey represented by layers 6, 4, 2 of the LGN¨ Dot: blind spot

Laminar structure of LGN Blind spot

15(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 16: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Interaction Potentials

Total free energy

1st term: retinotopy-generating term 2nd term: Correlational energy 3rd term: Vertical positioning energy

16(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

( , ) ( )corri i i i i i iE E e x E E y (5.7)

Page 17: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Correlational Energy Terms

Correlational energy for a type of interaction for a given terminal i

¨ dij: distance between terminals i and j¨ Wa: the set of all terminals participating in interaction a¨ Ba: the overall strength of the interaction energy of type a¨ : gradient ¨ ki: project number (?)

17(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

( )a

a ai ij

j W

E G d

(5.8)2

( ) exp( )( )ija a

ij ai

dG d B

s k

(5.9)

( ) 0.0015 0.4i ik k (5.10)

( )ik

Page 18: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Correlational Energy Terms

18(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 19: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Vertical positioning energy

¨ yi: the vertical position of the ith terminal

¨ a = 1.5¨ Kg : sequence of progressively

more negative constants for the six terminal types.

19(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

2( )i i i g iE y ay K y (5.10)

Page 20: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Induction of the Laminar Transition

20(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 21: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Trapping of the Transition by the Blind Spot Blind spot modeling

¨ Ghost magnocellular (layer 1) and prvocellular, ON po-larity (layer 6) terminals in a small portion of the con-tralateral eye

By simulated annealing

21(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 22: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

Summary

The Multiple Constraint Model Morphogenesis of the Lateral Geniculate Nucleus

22(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/

Page 23: Ch 5. The Patterning of Neural Connections 5.5 ~ 5.6 Adaptive Cooperative Systems, Martin Beckerman, 1997. Summarized by Kwonill, Kim Biointelligence Laboratory,

QnA

What can we infer from these models?

23(C) 2009, SNU Biointelligence Lab, http://bi.snu.ac.kr/