the dendrites of ca2 and ca1 pyramidal neurons ...v srv...
TRANSCRIPT
![Page 1: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/1.jpg)
Accepted manuscripts are peer-reviewed but have not been through the copyediting, formatting, or proofreadingprocess.
Copyright © 2017 the authors
This Accepted Manuscript has not been copyedited and formatted. The final version may differ from this version.
Research Articles: Cellular/Molecular
The dendrites of CA2 and CA1 pyramidal neurons differentially regulateinformation flow in the cortico-hippocampal circuit
Kalyan V. Srinivas1,3, Eric W. Buss1,5, Qian Sun1, Bina Santoro1, Hiroto Takahashi1, Daniel A. Nicholson5 and
Steven A. Siegelbaum1,2,3,4
1Department of Neuroscience2Department of Pharmacology3Howard Hughes Medical Institute4Kavli Institute for Brain Science Columbia University, 1051 Riverside Drive, New York, NY 10032, USA5Department of Neuroscience Rush University Medical Center, Chicago, IL 60612
DOI: 10.1523/JNEUROSCI.2219-16.2017
Received: 12 July 2016
Revised: 27 December 2016
Accepted: 7 February 2017
Published: 17 February 2017
Author contributions: K.V.S., D.A.N., and S.A.S. designed research; K.V.S., E.W.B., Q.S., B.S., and H.T.performed research; K.V.S., E.W.B., Q.S., and B.S. analyzed data; K.V.S., B.S., D.A.N., and S.A.S. wrote thepaper.
Conflict of Interest: The authors declare no competing financial interests.
This work was supported by the Howard Hughes Medical Institute. We thank Yeonsook Shin and Austen Sitkofor their help with biocytin stainings and the Neurolucida system. We also thank Anastasia Barnett for the helpwith immunohistochemistry.
Corresponding Authors: Kalyan V. Srinivas ([email protected]) or Steven A. Siegelbaum([email protected]) Postal Address: Columbia University, 1051 Riverside Drive, New York, NY 10032,USA.
Cite as: J. Neurosci ; 10.1523/JNEUROSCI.2219-16.2017
Alerts: Sign up at www.jneurosci.org/cgi/alerts to receive customized email alerts when the fully formattedversion of this article is published.
![Page 2: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/2.jpg)
5Department of Neuroscience Rush University Medical Center, Chicago, IL 60612
![Page 3: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/3.jpg)
Ih
![Page 4: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/4.jpg)
![Page 5: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/5.jpg)
![Page 6: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/6.jpg)
Ih
μ
°
![Page 7: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/7.jpg)
°
° Ω
μ
Ω
°
μ μ
![Page 8: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/8.jpg)
![Page 9: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/9.jpg)
N n
Dr Sl Sd
N =nπ Dr + Sl − Dr + Sd
θπ Dr + Sl − Dr + Sl θ Dr + Sd
θ =Dr + SdDr + Sl
![Page 10: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/10.jpg)
![Page 11: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/11.jpg)
R Ω C μ R Ω
Rm
C
SpineScale =SAshaft + SAspine
SAshaft
=πd + NA
πd
![Page 12: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/12.jpg)
SA μ SA
μ d
N A
μ
Rinput
Rinput
Ih Gh
Ih
Gh
Gh
Ih Ih
Gh
![Page 13: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/13.jpg)
Ih
τ τ
gh =g
+V +
Ih = gh V +
![Page 14: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/14.jpg)
Io Ko α
z F
R T g
g2
Io = g(V- Vrev) V Vrev
μ
![Page 15: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/15.jpg)
Amigo2-Cre
![Page 16: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/16.jpg)
μ μ μ
μ
![Page 17: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/17.jpg)
μ
Ih
![Page 18: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/18.jpg)
μ
μ μ
![Page 19: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/19.jpg)
μ
μ
μ
![Page 20: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/20.jpg)
Ih
Ih
Ih
Ih
Ih
![Page 21: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/21.jpg)
![Page 22: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/22.jpg)
Ih
Ih μ
Ih
μ
Ih
Ih
![Page 23: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/23.jpg)
Ih
Ih
Ih
![Page 24: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/24.jpg)
μ μ
μ μ
μ μ
μ μ
![Page 25: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/25.jpg)
μ
![Page 26: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/26.jpg)
μ
![Page 27: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/27.jpg)
Ih
Ih
Rm Ri
Cm
RInput
Rm Cm
RInput
Rm Ri
Ih Ih
![Page 28: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/28.jpg)
Ih
Ih
Ih
Ω (
μ
![Page 29: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/29.jpg)
![Page 30: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/30.jpg)
Ih
Ih
Ih
Ih
Ih
Ih Ih
Ih
μ
Ih Ih
![Page 31: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/31.jpg)
μ
μ
Ih
![Page 32: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/32.jpg)
Ih
![Page 33: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/33.jpg)
Ih
Ih
Ih
![Page 34: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/34.jpg)
Ih
Ih
Ih
Ih
Ih
Ih
![Page 35: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/35.jpg)
Ih
Ih
Ih
Ih
![Page 36: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/36.jpg)
Ih
![Page 37: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/37.jpg)
![Page 38: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/38.jpg)
![Page 39: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/39.jpg)
![Page 40: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/40.jpg)
![Page 41: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/41.jpg)
![Page 42: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/42.jpg)
![Page 43: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/43.jpg)
Ih
![Page 44: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/44.jpg)
![Page 45: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/45.jpg)
![Page 46: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/46.jpg)
![Page 47: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/47.jpg)
![Page 48: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/48.jpg)
![Page 49: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/49.jpg)
μ
μ
μ μ
![Page 50: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/50.jpg)
μ
Ih
![Page 51: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/51.jpg)
Rinput
μ
Ih
μ
μ
![Page 52: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/52.jpg)
μ
Ih
Ih
Ih
Rinput
![Page 53: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/53.jpg)
Ih
Ih
μ μ
![Page 54: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/54.jpg)
f
f μ f
μ
![Page 55: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/55.jpg)
![Page 56: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/56.jpg)
![Page 57: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/57.jpg)
![Page 58: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/58.jpg)
![Page 59: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/59.jpg)
![Page 60: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/60.jpg)
![Page 61: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/61.jpg)
![Page 62: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/62.jpg)
![Page 63: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/63.jpg)
![Page 64: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/64.jpg)
![Page 65: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/65.jpg)
![Page 66: The dendrites of CA2 and CA1 pyramidal neurons ...v srv txhvwlrqxvlqjdpxowlglvflsolqdu\dssurdfkrizkrohfhoovrpdw lfdqgghqgulwlfsdwfkfodps srw uhfruglqjv wzr skrwrq joxwdpdwh xqfdjlqj](https://reader033.vdocument.in/reader033/viewer/2022041722/5e4fa4c3cd2f2b7e3f13d526/html5/thumbnails/66.jpg)