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* indicates new v2.0 neuron type Name Supertype (ID:Family:Type:Subregion) CA1 (i)0313p-Sub_001 Hippocampo-subicular Projecting ENK+ S131504:Collateral-related:RO-targeting:CA1 Name derivation The name borrows from one of the cited descriptors in the literature that describes the projecting nature of the cell and its enkephalin-positivity. CA1 clarifies the subregion of origin, since the name is otherwise generic. Synonym(s) CA1 ENK-expressing cell CA1 ENK-expressing hippocampo-subicular cell CA1 ENK-expressing interneuron CA1 ENK-expressing projection neuron CA1 ENK-immunopositive cell CA1 enkephalin-expressing cell List of articles Click here to view the list Fuentealba P  (2008) J Neurosci Rhythmically active enkephalin-expressing GABAergic cells in the CA1 area of the hippocampus project to the subiculum and preferentially innervate interneurons. Fuentealba P, Tomioka R, Dalezios Y, Marton LF, Studer M, Rockland K, Klausberger T, Somogyi P. J Neurosci, 2008 Oct 1, 28 (40), Pages: 10017 - 10022 PMID: 18829959; DOI: 10.1523/JNEUROSCI.2052-08.2008 Tags: morphology, marker, connectivity, phases, synapse probabilities

Morphology

Soma
	CA1:SR

Axons
	CA1:SR
	CA1:SP
	CA1:SO
	SUB:PL

Dendrites
	CA1:SR
	CA1:SO

Representative figure

Rhythmically active enkephalin-expressing GABAergic cells in the CA1 area of the hippocampus project to the subiculum and preferentially innervate interneurons. Fuentealba P, Tomioka R, Dalezios Y, Marton LF, Studer M, Rockland K, Klausberger T, Somogyi P J Neurosci, 2008 Oct 1, 28 (40), pages: 10017 - 10022 PMID: 18829959; DOI: 10.1523/JNEUROSCI.2052-08.2008

"Figure 2. Subicular projection and synaptic output of an identified ENK[enkephalins]-expressing interneuron {CA1 Hippocampo-subicular Projecting ENK+} [in CA1] (P30b) recorded in vivo. ... C, Reconstruction of the cell. The dendritic arbor (orange) is complete; the axon (yellow) is presented partially in three panels from rostral (section 86) to caudal (section 63) levels. The axon formed a columnar arbor near the soma and an elongated plexus (2.5mm) in the alveus, continuing into and innervating the subiculum. Four light ellipses depict the location of some synaptically targeted interneurons."

Molecular markers

Positive
	CoupTF II
	ENK
	mGluR1
	mGluR1a
	vGAT

Negative
	CB1 (confirmed by inference)
	CCK (confirmed by inference)
	ChAT
	CR
	GABAa\alpha 6 (inference)
	Gaba-a-alpha
	GlyT2
	Mus2R
	NPY (inference)
	PV (confirmed by inference)
	Sub P Rec
	vAChT
	vGluT3 (multiple confirming inferences)
	VIP

Mixed expression
	SOM (unresolved inferential conflict)

Electrophysiological properties

None known

Notes

- inhibitory because GABAergic (Fuentealba et al., 2008) - interneuron specific: - 57% of CA1 targets and 100% of SUB targets were interneurons (Fuentealba 2008) - up to half the targets are other interneurons (Somogyi 2010) - 11.3% (7/62) of hippocampo-subicular nonpyramidal projection cells were ENK-immunopositive - Fuentealba paper refers to these cells as a "population," though only one cell could be positively identified

Firing Pattern

Images
	No image is associated with this neuron

Parameters
	No parameter associated with this neuron

ModelDB Model

Model Accession Number

PubMed ID

Sources of Input

Known sources Excitatory or Inhibitory none known

Potential sources Excitatory or Inhibitory CA3 Pyramidal CA3c Pyramidal CA3c Umbrella Pyramidal* CA3 Trilaminar CA2 Pyramidal CA2 Bistratified CA1 Pyramidal CA1 Deep Pyramidal CA1 Superficial Pyramidal CA1 Radiatum Giant CA1 Back-Projection CA1 Bistratified CA1 Hippocampo-subicular Projecting ENK+ CA1 Ivy CA1 Interneuron Specific LM-R CA1 Interneuron Specific LMO-O CA1 LMR CA1 Interneuron Specific LMR-O* CA1 LMR-R* CA1 Interneuron Specific LMR-R CA1 LMR Projecting CA1 Recurrent O-LM CA1 O-LMR CA1 Recurrent O-LMR* CA1 Interneuron Specific O-R CA1 O-RP* CA1 Interneuron Specific O-Targeting QuadD CA1 Oriens/Alveus CA1 Oriens-Bistratified CA1 Oriens-QuadA Projecting CA1 O-RPO* CA1 QuadD-RP* CA1 Quadrilaminar CA1 Interneuron Specific R-O CA1 R-Receiving Apical-Targeting CA1 R-RO* CA1 R-RP* CA1 Radiatum CA1 Interneuron Specific RO-O CA1 Interneuron Specific RO-R* CA1 RO-RP* CA1 RP-RPO* CA1 Schaffer Collateral-Associated CA1 Schaffer Collateral-Correlated* CA1 Schaffer Collateral-Receiving R-Targeting CA1 SO-SO CA1 SO-SO Projecting* CA1 Trilaminar CA1 Radial Trilaminar Sub CA1-Projecting Pyramidal EC LIII Small Pyramidal EC LIII Pyramidal

Potential sources known to be avoided Excitatory or Inhibitory none known

Targets of Output

Known targets Excitatory or Inhibitory CA1 Pyramidal

Potential targets Excitatory or Inhibitory CA2 Wide-Arbor Basket CA1 Deep Pyramidal CA1 Superficial Pyramidal CA1 Radiatum Giant CA1 Axo-axonic CA1 Horizontal Axo-axonic CA1 Back-Projection CA1 Basket CA1 Horizontal Basket CA1 R-P Basket* CA1 Trilayer Basket* CA1 Vertical Basket* CA1 Basket CCK+ CA1 Horizontal Basket CCK+* CA1 Trilayer Basket CCK+* CA1 Bistratified CA1 RO-receiving CA3-targeting* CA1 Hippocampo-subicular Projecting ENK+ CA1 Ivy CA1 Interneuron Specific LMO-O CA1 LMR CA1 LMR-LM* CA1 Interneuron Specific LMR-O* CA1 LMR-R* CA1 Interneuron Specific LMR-R CA1 LMR Projecting CA1 O-LM CA1 Recurrent O-LM CA1 O-LMR CA1 Recurrent O-LMR* CA1 Interneuron Specific O-R CA1 O-RP* CA1 Interneuron Specific O-Targeting QuadD CA1 Oriens/Alveus CA1 Oriens-Bistratified CA1 Oriens-QuadA Projecting CA1 OR-LM CA1 O-RPO* CA1 Perforant Path-Associated CA1 Perforant Path-Associated QuadD CA1 QuadD-RP* CA1 Quadrilaminar CA1 Interneuron Specific R-O CA1 R-Receiving Apical-Targeting CA1 R-RO* CA1 R-RP* CA1 Radiatum CA1 Interneuron Specific RO-O CA1 Interneuron Specific RO-R* CA1 RO-RP* CA1 RP-RPO* CA1 Schaffer Collateral-Associated CA1 Schaffer Collateral-Correlated* CA1 Schaffer Collateral-Receiving R-Targeting CA1 SO-SO CA1 SO-SO Projecting* CA1 Trilaminar CA1 Radial Trilaminar Sub Wide-arbor Polymorphic* Sub Recurrent Polymorphic-Moleculare*

Potential targets known to be avoided Excitatory or Inhibitory none known

Oscillation Phase Locking

Category	Value
Theta (deg):	62.00
Theta range:	[62.00, 62.00]
Theta measurements:	1
Theta representative selection:	rats, adult, male, juxtacellular, urethane plus supplemental doses of ketamine and xylazine
SWR ratio:	0.09
SWR ratio range:	[0.09, 0.09]
SWR ratio measurements:	1
SWR ratio representative selection:	rats, adult, male, juxtacellular, urethane plus supplemental doses of ketamine and xylazine
In vivo firing rate (Hz):	5.50
Firing rate range:	[5.50, 5.50]
Firing rate measurements:	1
Firing rate representative selection:	rats, adult, male, juxtacellular, urethane plus supplemental doses of ketamine and xylazine
Other:	gamma and 1 other
Other representative selection:	rats, adult, male, juxtacellular, urethane plus supplemental doses of ketamine and xylazine