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. Author manuscript; available in PMC: 2017 Feb 23.
Published in final edited form as: J Comp Neurol. 2016 May 5;524(14):2803–2827. doi: 10.1002/cne.23992

Table 2.

c-Fos-immunoreactivity (Fos-ir) in brains from fasted and refed rats mapped to Swanson (2004)

Cell Group or Region (Abbreviation, Swanson Atlas Level) Density of Fos-ir(1)

Fasted Refed
Cerebrum (CH)
 Cerebral Nuclei
  Pallidum
   Globus pallidus (GP)
    Medial globus pallidus (GPi, 27) (+) (+)
    Lateral globus pallidus (GPe, 27) (−) (−)
   Substantia innominata (SI, 27) (−) (−)
   Medial septal complex (MSC)
    Medial septal nucleus (MS, 18) ++++ +++*
   Bed nuclei of terminal stria (BST)
    Anteromedial area (BSTam, 18) +++ ++++*
  Striatum
   Lateral septal nucleus (LS)
    rostral (rostroventral) part (LSr, 18) + +*
    ventral part (LSv, 18) ++++ +++
   Caudoputamen (CP, 27) (−) (−)
     (CP, 30) (−) (+)
   Central amygdalar nucleus (CEA)
    Medial part (CEAm, 27) (−) (+++)
    Lateral part (CEAl, 27) (−) (++)
     (CEAl, 30) (−) (−)
   Intercalated amygdalar nuclei (IA, 27) (−) (−)
     (IA, 30) (−) (−)
   Medial amygdalar nucleus (MEA)
    Anteroventral part (MEAav, 27) (+) (++)
    Anterodorsal part (MEAad, 27) (+) (+)
    Posterodorsal part, sublayer c (MEApd.c, 30) (−) (++)
 Cerebral Cortex (CTX)
  Cortical Plate (CTXpl)
   Limbic region
    Olfactory region (OLF)
     Piriform area (PIR)
      molecular layer (PIR1, 27) (−) (+)
       (PIR1, 30) (−) (+)
      pyramidal layer (PIR2, 27) (−) (−)
       (PIR2, 30) (−) (+)
      polymorph layer (PIR3, 27) (−) (−)
       (PIR3, 30) (−) (++)
     Cortical amygdalar complex
      Cortical amygdalar area (COA)
       Anterior part (COAa, 27) (+) (++)
       Posterior part (COAp, 30)
        Medial zone (COApm, 30)
         layer 1 (−) (+)
         layer 3 (+) (+)
        Lateral zone (COApl, 30)
         layer 1 (−) (−)
         layer 2 (−) (+)
         layer 3 (+) (+)
      Piriform-amygdalar area (PAA, 30)
       layer 1 (−) (+++)
       layer 3 (−) (−)
    Retrohippocampal region (RHP)
     Entorhinal area (ENT)
      Lateral part (ENTl)
       layer 1 (ENTl1) (−) (−)
       layer 2 (ENTl2) (−) (−)
       layer 3 (ENTl3) (−) (−)
       layer 4 (ENTl4) (−) (−)
       layer 5 (ENTl5) (−) (−)
       layer 6 (ENTl6) (−) (−)
    Cingulate region (CNG)
     Prelimbic area (PL)
      layer 1 (PL1, 8) - +*
      layer 2 (PL2, 8) - ++*
      layer 3 (PL3, 8) - +*
      layer 5 (PL5, 8) - ++++*
      layer 6a (PL6a, 8) - +++*
     Anterior cingulate area, dorsal part (ACAd)
      layer 1 (ACAd1, 8) - +*
      layer 5 (ACAd5, 8) + +
      layer 6a (ACAd6a, 8) + +
      layer 6b (ACAd6b, 8) - -
    Frontal region (FRO)
     Somatomotor areas (MO)
      Secondary somatomotor areas (MOs)
       layer 1 (MOs1, 8) - +*
       layers 2/3 (MOs2/3, 8) - +*
       layer 5 (MOs5, 8) + +
       layer 6a (MOs6a, 8) ++* +
       layer 6b (MOs6b, 8) - -
    Temporal region (TE)
     Perirhinal area (PERI)
      layer 1 (PERI1) (−) (−)
      layer 3 (PERI3) (−) (−)
      layer 5 (PERI5) (+) (−)
     Ectorhinal area (ECT)
      layer 1 (ECT1) (−) (+)
      layer 3 (ECT3) (−) (+)
      layer 5 (ECT6) (−) (+)
     Temporal association areas (TEa)
      layer 1 (TEa1) (−) (−)
      layer 3 (TEa2) (−) (−)
      layer 4 (TEa4) (−) (+)
      layer 5 (TEa5) (−) (+)
    Cortical subplate (CTXsp)
     Basolateral amygdalar complex
      Basomedial amygdalar nucleus (BMA)
       Anterior part (BMAa, 27) (−) (+)
       Posterior part (BMAp, 27) (2) (−) (−)
        (BMAp, 30) (−) (+)
       Basolateral amygdalar nucleus (BLA, 30)
        Anterior part (BLAa, 27) (−) (−)
        Posterior part (BLAp, 30) (−) (−)
        Lateral amygdalar nucleus (LA, 27) (−) (−)
         (LA, 30) (−) (−)
        Posterior amygdalar nucleus (PA, 30) (+) (−)
         (PA, 33) - -
      Endopiriform nucleus (EP)
       Ventral part (EPv, 27) (−) (−)
        (EPv, 30) (−) (−)
       Dorsal part (EPd, 27) (−) (−)
        (EPd, 30) (−) (−)
Interbrain (IB)
 Hypothalamus (HY)
  Periventricular hypothalamic zone
   Terminal lamina (lam)
    Median preoptic nucleus (MEPO, 18) + +
   Internuclear area, hypothalamic periventricular part (I, 30) ++* +
   Anterodorsal preoptic nucleus (ADP, 18) ++++ ++++
   Medial preoptic area (MPO, 18) ++* +++
   Parastrial nucleus (PS, 18) ++ ++++*
   Subparaventricular zone (SBPV, 26) +* +
     (SBPV, 27) +* +
   Paraventricular hypothalamic nucleus, magnocellular division (PVHm)
    Posterior magnocellular part (PVHpm)
     lateral zone (PVHpml, 26) + +++*
   Paraventricular hypothalamic nucleus, parvicellular division (PVHp)
    Periventricular part (PVHpv, 27) + +
    Medial parvicellular part, dorsal zone (PVHmpd, 27) + !!!!
    [Lateral parvicellular part] (PVHlp, 27) ++++ !!!!
    [Forniceal part] (PVHf, 27) +* ++++
   Supraoptic nucleus (SOg)
    retrochiasmatic part (SOr) - -
   Neurohypophysis
    Infundibulum (INF)
     Median eminence (ME, 30) - +
   Arcuate hypothalamic nucleus (ARH, 30) - ++++
   Periventricular hypothalamic nucleus intermediate part (PVi) - ++
   Dorsomedial hypothalamic nucleus (DMH)
    Anterior part (DMHa, 30) + +
    Posterior part (DMHp, 30) + ++
    Ventral part (DMHv, 30) ++* !!!!!
   Posterior hypothalamic nucleus (PH, 30) ++* +*
    (PH, 33) !!!! !!!!
   Periventricular hypothalamic nucleus, posterior part (PVp, 33) +++* ++
  Medial hypothalamic zone
   Anterior hypothalamic nucleus (AHN)
    Central part (AHNc, 27) +* ++*
    Posterior part (AHNp, 27) ++* +++
    Dorsal part (AHNd, 27) - +
   Ventromedial hypothalamic nucleus (VMH)
    Ventrolateral part (VMHvl, 27) + +
     (VMHvl, 30) ++* ++
    Central part (VMHc, 27) +* ++*
     (VMHc, 30) +* ++
    Dorsomedial part (VMHdm, 27) +* +*
   Ventral premammillary nucleus (PMv, 33) +++* -
   Dorsal premammillary nucleus (PMd, 33) + ++*
    Mammillary body (MB)
     Tuberomammillary nucleus (TM)
      ventral part (TMv, 33) + +
     Medial mammillary nucleus (MMg)
      median part (MMe, 33) - -
     Supramammillary nucleus (SUM)
      lateral part (SUMl, 33) +++* +
  Lateral hypothalamic zone
   Lateral preoptic area (LPO, 18) +* ++++
   Lateral hypothalamic area (LHA)
    Anterior group (LHAag)
     Anterior region (LHAa)
      intermediate zone (LHAai, 26) +++ ++++
      dorsal zone (LHAad, 26) + ++++
    Middle group (LHAmg)
     Medial tier (LHAm)
      Juxtaventromedial region (LHAjv)
       Ventral zone (LHAjvv, 27) (3) ++ ++
        (LHAjvv, 30) ++++* +++
       Dorsal zone (LHAjvd, 27) (3) + ++*
      Juxtaparaventricular region (LHAjp, 26) + ++++
      Juxtadorsomedial region (LHAjd, 27) + +
       (LHAjd, 30) ++++* ++++
     Perifornical tier (LHApf)
      Subfornical region (LHAsf)
       Anterior zone (LHAsfa, 27) +++* !!!!*
       Posterior zone (LHAsfp, 30) ++ +++*
       Premammillary zone (LHAsfpm, 33) ++++ +++*
      Suprafornical region (LHAs, 30) ++++* ++
     Lateral tier (LHAl)
      Tuberal nucleus (TU)
       Subventromedial part (TUsv, 27) - ++++
        (TUsv, 33) - -
       Intermediate part (TUi, 27) (3) +* -
        (TUi, 33) - +
       Terete part (TUte, 33) - +
       Lateral part (TUl, 33) + -
      Ventral region (LHAv)
       Medial zone (LHAvm, 30) ++++* ++++*
       Lateral zone (LHAvl, 30) + +
        (LHAvl, 33) + +
       Parvicellular region (LHApc, 27) + +
       Magnocellular nucleus (LHAma, 30) + -
      Dorsal region (LHAd, 27) !!!! !!!!
       (LHAd, 30) ++++ ++++
    Posterior group (LHApg)
     Posterior region (LHAp, 33) !!!!* !!!!*
     Parasubthalamic nucleus (PSTN, 33) ++ !!!!*
     Subthalamic nucleus (STN, 30) - -
      (STN, 33) +++ !!!!
Thalamus (TH)
 Ventral part of thalamus (VNT)
  Zona incerta, general (ZIg)
   Zona incerta (ZI, 26) +++* ++++
    (ZI, 27) +++* !!!!!
    (ZI, 30) ++++ +++*
    (ZI, 33) +++* ++*
   Fields of Forel (FF, 33) ++++ +
 Dorsal part of thalamus (DOR)
  Midline thalamic nuclei (MTN)
   Nucleus reuniens (RE)
    Rostral division (REr, 26) (4) ++++* +++
    Caudal division (REca, 27) (4) ++ +++
   Paraventricular thalamic nucleus (PVT, 25) ++ ++++
   Paratenial nucleus (PT, 25) ++ ++++
  Anterior thalamic nuclei (ATN)
   Anteromedial thalamic nucleus (AM, 25)
    Dorsal part (AMd) + -
  Intralaminar thalamic nuclei (ILM)
   Central medial thalamic nucleus (CM, 25) + ++
    (CM, 33) + -
  Medial thalamic nuclei (MED)
   Perireuniens nucleus (PR, 27) +* +*
    (PR, 30) - +
   Submedial thalamic nucleus (SMT, 30) - -
   Mediodorsal nucleus of the thalamus (MD)
    Medial part (MDm, 25) - +*
  Ventral thalamic nuclei
   Ventral medial thalamic nucleus (VM, 30) - -
   Ventral posterior thalamic nucleus (VP)
    Ventral posteromedial thalamic nucleus, general (VPMg)
     Parvicellular part (VPMpc, 33) - +*
   Ventral posterolateral thalamic nucleus, general (VPLg)
     Parvicellular part (VPLpc, 33) - -
   Subparafascicular nucleus (SPF)
    Magnocellular part (SPFm, 33) +* ++*
Midbrain (MB)
 Tegmentum (TG)
  Interpeduncular nucleus (IPN)
   apical subnucleus (IPNa, 43) +++ +
   dorsomedial subnucleus (IPNd, 43) + ++
   lateral subnucleus (IPNl, 43)
    dorsal part (IPNld) - -
    intermediate part (IPNli) + -
    ventral part (IPNlv, 43) + -
   intermediate subnucleus (IPNi, 43) + -
   central subnucleus (IPNc, 43) + -
  Central linear raphe nucleus (CLI, 43) + -
  Superior central raphé nucleus (CS)
   medial part (CSm, 43) + -
  Dorsal raphe nucleus (DR, 43) + -
   (DR, 49) !!!! +++
  Ventral tegmental area (VTA, 43) - +
  Midbrain reticular nucleus (MRN)
   Magnocellular part (MRNm, 43) ++++* +++
   Parvicellular part (MRNp, 43) ++* -
   Retrorubral area (RR, 43) ++++* +
  Periaqueductal gray (PAG)
   Medial division (PAGm, 43) - +
   Ventrolateral division (PAGvl, 43) !!!! !!!!
   Dorsolateral division (PAGdl, 43) - -
   Dorsal division (PAGd, 43) ++ ++
  Midbrain nucleus of trigeminal nerve (MEV, 49) + ++
   (MEV, 50) + !!!!
   (MEV, 69) - -
  Nucleus sagulum (SAG) - +
  Parabigeminal nucleus (PBG, 43) - -
  Nucleus of brachium of inferior colliculus (NB) +++* !!!!
 Tectum (TC)
  Superior colliculus (SC)
   zonal layer (SCzo, 43) - +
   superficial gray layer (SCsg, 43) - -
   sublayer a of intermediate gray layer (SCig.a, 43) - ++*
   sublayer b of intermediate gray layer (SCig.b, 43) ++* ++++*
   sublayer c of intermediate gray layer (SCig.c, 43) ++* +++*
  Inferior colliculus (IC) - -
   Central nucleus (ICc, 49) !!!!!* -
    (ICc, 50) - -
   Dorsal nucleus (ICd, 49) !!!! -
    (ICd, 50) - -
   External nucleus (ICe, 43) +++* ++++
Rhombic Brain (RB)
 Hindbrain (HB)
  Pons (P)
   Motor nucleus of trigeminal nerve (V)
    Magnocellular part (Vma, 49) + !!!!*
     (Vma, 50) +++ !!!!!*
    Parvicellular part (Vpc, 49) +* -
     (Vpc, 50) - +
   Pontine nuclei, general (PGg)
    Pontine nuclei (PG, 43) !!!!! +
   Tegmental reticular nucleus (TRN, 49) (5) ++ +
    (TRN, 50) (5) +++ ++
   Magnus raphe nucleus (RM, 49) + +
    (RM, 50) + +
   Pontine raphe nucleus (RPO, 49) +++ +++*
    (RPO, 50) +++* +*
   Pontine reticular nucleus, Caudal part (PRNc, 49) !!!!* ++++
    (PRNc, 50) !!!!* ++++*
   Pedunculopontine nucleus (PPN, 43) +++ ++*
   Supratrigeminal nucleus (SUT, 49)Pontine central gray, general (PCGg)
    Pontine central gray (PCG, 49) !!!! ++++*
     (PCG, 50) +++* ++*
    Nucleus incertus (NI)
     compact part (NIc, 50) ++++ ++++
     diffuse part (NId, 50) ++++ ++*
    Lateral tegmental nucleus (LTN, 49) + +++*
    Dorsal tegmental nucleus (DTN, 49) +++* +
     (DTN, 50) !!!!* ++++*
    Sublaterodorsal nucleus (SLD, 49) (5) ++++* +
     (SLD, 50) ++* +
    Laterodorsal tegmental nucleus (LDT, 49) ++++ +++
     (LDT, 50) !!!!* +++*
    Barrington’s nucleus (B, 49) + +*
     (B, 50) ++++* !!!!*
    Locus ceruleus (LC, 49) - ++++
     (LC, 50) ++ ++++
   Parabrachial Nucleus (PB)
    Medial division (PBm)
     Ventral medial part (PBmv, 50) +* ++++*
     Medial medial part (PBmm, 49) + !!!!
      (PBmm, 50) + ++++*
    Kölliker-Fuse subnucleus (KF, 49) +* !!!!!*
    Lateral division
     Ventral lateral part (PBlv, 49) + !!!!*
      (PBlv, 50) - !!!!*
     Central lateral part (PBlc, 49) ++ !!!!*
      (PBlc, 50) ++* !!!!!*
     External lateral part (PBle, 49) - ++*
      (PBle, 50) - !!!!*
     Dorsal lateral part (PBld, 49) +* +++*
   Principal sensory nucleus of trigeminal nerve (PSV, 49) !!!!* ++++*
    (PSV, 50) !!!!* +++*
   Nucleus of lateral lemniscus (NLL)
    ventral part (NLLv, 43) + +
   Superior olivary complex (SOC)
    medial part (SOCm, 49) (5) +* -
     (SOCm, 50) (5) + +
    lateral part (SOCl, 50) (5) ++* +++*
    periolivary region (POR, 49) (5) !!!!* !!!!*
     (POR, 50) (5) !!!!* !!!!*
   Nucleus of trapezoid body (NTB, 49) + !!!!
    (NTB, 50) + +++*
  Medulla (MY)
   Hypoglossal nucleus (XII, 69) - +
   Ambiguus nucleus, dorsal division (AMBd, 69) - -
   Ambiguus nucleus, ventral division (AMDv, 69) - +
   Dorsal motor nucleus of vagus nerve (DMX, 69) - +*
   Obscurus raphe nucleus (RO, 69) - -
   Pallidal raphe nucleus (RPA) + -
   Parvicellular reticular nucleus (PARN, 69) + +
   Medullary reticular nucleus (MDRN)
    Ventral part (MDRNv, 69) + +*
   Inferior olivary complex (IO)
    Medial accessory olive (IOma, 69) + +
   Lateral reticular nucleus (LRN, 69)
    Magnocellular part (LRNm) ++ +
    Cochlear nuclei (CN)
     Ventral cochlear nucleus, anterior part (VCOa, 49) !!!!* ++*
      (VCOa, 50) ++* +
     Subpeduncular granular region (CNspg, 49) - -
      (CNspg, 50) !!!!* ++*
    Parvicellular part (LRNp) - +
   Paramedian reticular nucleus (PMR, 69) - -
   Parasolitary nucleus (PAS, 69) - +*
   Nucleus of solitary tract (NTS)
    Medial part (NTSm)
     Caudal zone (NTSm, 69) - !!!!!
    Gelatinous part (NTSge, 69) - +++*
    Lateral part (NTSl, 69) - ++*
    Commissural part (NTSco, 69) - ++++*
   Area postrema (AP, 69) ++++ !!!!
   Dorsal column nuclei (DCN, 69)
    Gracile nucleus, general (GRg)
     gracile nucleus, principal part (GR) + +
    Cuneate nucleus (CU, 69) - -
    External cuneate nucleus (ECU, 69) ++ ++
   Paratrigeminal nucleus (PAT, 69) - +++*
   Spinal nucleus of the trigeminal nerve (SPV)
    Interpolar part (SPVI, 69) + ++*
    Caudal part (SPVC, 69) - +

Density of Fos-ir(1)

Fos Scattered in White Matter Tracts (Abbreviation, Atlas Level) Fasted Refed

Cerebrospinal Axis
Spinal tract of the trigeminal nerve (sptV, 69) ++ +++
Tegmentum
Brachium of inferior colliculus (bic, 43) +++* !!!!
Pons
Middle cerebellar peduncle (mcp, 43) +++ -
Ventral spinocerebellar tract (sctv, 49) !!!! !!!!*
  (sctv, 50) !!!!* +++
Pyramid (py, 49) ++ !!!!!
  (py, 50) +++ !!!!*

This tabular, hierarchical arrangement for anatomical regions is based (with minor variations) on the provisional topographic histological groupings provided in Table C, Rat CNS Gray Matter Regionalization, BM4 Beta 3.1 and Table D, Rat CNS White Matter Tracts, BM4 Beta 1, available at the time of this writing in the online material from Swanson LW, 2015a (white papers dated 6/25/15 for Table C and 4/5/15 for Table D). This order standardizes topographic ordering based on embryological/developmental sequence, with rostral before caudal, then medial before lateral, and then ventral before dorsal. The names of the structures have evolved from Swanson (2004) to become standardized by Swanson (2015a). Therefore, all newer, standard terms for each brain part are used whenever possible (these are denoted by underlining; for example, ‘paratrigeminal nucleus’). If the older term from Swanson (2004) is retained, it is denoted in brackets (e.g., ‘[Lateral parvicellular part]’ of the paraventricular hypothalamic nucleus). Note, however, that all symbols of the structures themselves (e.g., ‘GPi’) are from Swanson LW (2004) and correspond to the symbols within the maps presented in Figure 2.

Explanations of symbols

Scale (c-Fos-IR cells): 0: – | 1–5 cells: + | 6–10 cells: ++ | 11–20 cells: +++ | 20–30 cells: ++++ | 31–99 cells: !!!! | >100 cells: !!!!!;

Other superscripted notations:

(1)

The scoring represents combined bilateral counts at each mapped level (i.e., the counts were performed on the maps shown in Figure 2, not on the tissue itself). Asterisks (*) denote a bilateral difference in the same subject (i.e., the combined bilateral count reflects mainly signal on one side, as it was not distributed uniformly between sides). Scoring symbols marked in parentheses (e.g., ‘(+++)’) represent unilateral counts, since in some cases, only one side of the tissue shared a common plane of section and therefore was the only side mapped.

(2)

For Level 27, the boundaries for the BMAp were indistinct in Nissl-stained tissue analyzed for the refed cohort, and were inferred (this uncertainty is denoted by pale yellow colored boundaries; see Figure 2, Panel D4).

(3)

For Level 27, the boundaries for the LHAjvd, LHAjvv, and TUi were inferred, as they were indistinct in Nissl-stained sections (this uncertainty is denoted by pale yellow colored boundaries; see Figure 2, Panels D2, E1 and E2).

(4)

For the reuniens nucleus, boundaries for the subdivisions were inferred, as they were indistinct in the Nissl-stained sections (this uncertainty is denoted by pale yellow colored boundaries; see Figure 2, Panels C1, C2 and D2). Therefore, c-Fos+ counts represent those found in the structure’s main divisions (rostral vs. caudal) rather than from specific subdivisions.

(5)

For Levels 49 and 50, the boundaries for the SLD, TRN, SOCm, SOCl, and POR were often indistinct in Nissl-stained tissue analyzed for one or both cohorts, and were therefore inferred (this uncertainty is denoted by pale yellow colored boundaries; see Figure 2, Panels H1–H4).