Table 2.
Characteristics of zonal function in (a) rat (b) mouse, and (c) human adult adrenal glands.
| (a) Rat | ||||
|---|---|---|---|---|
| Factor/Gene | Glomerulosa | Fasciculata/ | Reticularis | References |
| STEROIDOGENESIS | ||||
| CYP11A | +/−? | +++ | Roskelley and Auersperg, 1990; Halder et al., 1998; Mitani et al., 1999; Guasti et al., 2013b; but see Engeland and Levay-Young (1999) | |
| CYP11B1 | − | +++ | +/− | Ho and Vinson, 1993; Halder et al., 1998; Mitani et al., 1999 |
| CYP11B2 | + | − | Halder et al., 1998; Peters et al., 1998; Mitani et al., 1999 | |
| CYP21 | +/− | +++ | Chang et al., 2011, 2013 | |
| 3β-HSD | + | +++ | Dupont et al., 1991; Engeland and Levay-Young, 1999; Pignatelli et al., 1999 | |
| 11β-HSD1 | − | − | + | Shimojo et al., 1996 |
| 11β-HSD2 | − | ++ | ++ | Smith et al., 1997; Atanassova and Koeva, 2012 |
| StAR | − | +++ | Peters et al., 1998, 2007 | |
| RECEPTORS | ||||
| ATR1 | +++ | − | − | Lehoux et al., 1997 |
| MC2R | +/− Inner ZU, | +++ | +/− | Gorrigan et al., 2011 |
| MRAP | +/− see text | +++ | +/− | |
| MRAP2 | =/− | +/− | +/− | |
| MC5R | +++ | + | Griffon et al., 1994; van der Kraan et al., 1998 | |
| ZONAL HOMEOSTASIS | ||||
| Shh | +++ | − | Guasti et al., 2011, 2013b | |
| Pref-1/dlk1 | +++ | − | Halder et al., 1998; Raza et al., 1998; Whitworth and Vinson, 2000 | |
| NEUROTRANSMITTER | ||||
| VIP | + | − | Holzwarth et al., 1987; Bernet et al., 1994 | |
| NPY | + | − | Holzwarth et al., 1987; Bernet et al., 1994; Renshaw and Hinson, 2001 | |
| Catecholamines | + | − | Holzwarth et al., 1987; Bornstein and Ehrhart-Bornstein, 1992 | |
| CYTOKINES | ||||
| IGIF | − | +++ | ++ | Conti et al., 1997 |
| *IL-6 | + | Judd and MacLeod, 1992 | ||
| *TNF | + | Judd and MacLeod, 1995 | ||
| GROWTH FACTORS | ||||
| Prorenin | + | ++ | + | Ho and Vinson, 1998 |
| bFGF | + | + | Chambaz et al., 1994; Ho and Vinson, 1995, 1997 | |
| IGF | + | + | ||
| TGFβ | − | − | ||
| SIGNALING | ||||
| ANP | +++ | − | Lai et al., 2000 | |
| AgRP | − | +++ | +++ | Bicknell et al., 2000 |
| Prorenin | − | + | Ho and Vinson, 1998 | |
| MAPK:ERK-1, ERK-2 | ++ | − | McNeill et al., 1998; McNeill and Vinson, 2000; Vinson et al., 2000; Guasti et al., 2013b | |
| SF-1 | ++ | ++ | Raza et al., 1998 | |
| Dab2 | +++ | − | Romero et al., 2007 | |
| Nek 2b | + | +++ | − | de Mendonca et al., 2014 |
| Notch1 | +++ | ++ | + | |
| Notch2 | +capsule | − | − | |
| Notch3 | +++ | ++ | + | |
| DETOXIFICATION | ||||
| **MnSOD | − | ++ | Raza and Vinson, 2000 | |
| CLOCK GENES | ||||
| PER 1 | (?) | +++ (reticularis) | Fahrenkrug et al., 2008 | |
| PER 2 Bmal 1 | ||||
| OTHER | ||||
| IZA | − | +++ | ++ | Barker et al., 1992; Halder et al., 1998; Vendeira et al., 1999; Raza et al., 2001 |
| OAT1 | − | ++ | Beery et al., 2003 | |
| Oatp1,2 | − | ++ | ||
| Oatp3 | ++ | − | ||
| AsP | +++ | + | Bicknell et al., 2001 | |
| EphA2, EphA2, EphA3 | ++ | − | Brennan et al., 2008 | |
| Spexin | +++ | ++ | ++ | Rucinski et al., 2010 |
| *Release from incubated glomerulosa cells **immunoblotting | ||||
| (b) Mouse | ||||
| Factor/Gene | Glomerulosa | Fasciculata/Reticularis | X-Zone | References |
| STEROIDOGENESIS | ||||
| CYP11B1 | − | +++ | − (?) | Mukai et al., 2003; King et al., 2009 |
| CYP11B2 | +++ | − | ||
| 20α-HSD | − | − | ++ | Hershkovitz et al., 2007 |
| 3β-HSD | +++ | +++ | − | |
| CYP21 | +/− | +++ | Chang et al., 2013 | |
| RECEPTORS | ||||
| ATR1 | +++ | − | − | Huang et al., 2013 |
| SIGNALING | ||||
| Shh | +++ | − | King et al., 2009 | |
| Dax1 | +++ | − | Kim et al., 2009 | |
| Mukai et al., 2002; Scheys et al., 2011 | ||||
| GATA4/GATA6 GATA4 in outer capsule only | +++ | |||
| β-catenin | +++ | +/− | Tevosian et al., 2015 | |
| Pihlajoki et al., 2015 | ||||
| CLOCK GENES | ||||
| Bmal1, CLOCK, Cry2 | +++ | ++ | Oster et al., 2006 | |
| Per1, Per2, Per3, Rev-Erbα | ||||
| Pik3c2g | − | − | +++ | Pihlajoki et al., 2013 |
| DETOXIFICATION | ||||
| AKR1B7 | −? | ++ | Aigueperse et al., 2001; Martinez et al., 2001 | |
| Grx 1 | +/− | +++ | + | Godoy et al., 2011 |
| Grx 2,5 | ++ | + | ++ | |
| Prx 1,4,5 | + | +++ | + | |
| Prx 3 | + | + | ++ | |
| Trx 1,2 | + | +++ | + | |
| TrxR2 | + | − | + | |
| Other | ||||
| AZ-1 | +++ | ++(?) | ++(?) | Mukai et al., 2003; Bastida et al., 2007; Li et al., 2007 |
| ODC | +++ | ++ | − | |
| (c) Human | ||||
| Factor/Gene | Glomerulosa | Fasciculata | Reticularis | References |
| STEROIDOGENESIS | ||||
| CYP11B1 | − | +++ | +++ | Nishimoto et al., 2010 |
| CYP11B2 | + | − | − | |
| †CYP11A | +++ | +++ | Rege et al., 2014 | |
| CYP17 | +++ | +++ | Nishimoto et al., 2010 | |
| CYPB5 | + | +++ | Rainey et al., 2002 | |
| SULT2A1 | + | +++ | Rainey and Nakamura, 2008 | |
| 3β-HSD | + | +++ | +/− | Nishimoto et al., 2010; but see Rainey et al. (2002) |
| †3β-HSD2 | +++ | + | Rege et al., 2014 | |
| †AKR1C3 | + | |||
| ZONAL HOMEOSTASIS | ||||
| DACH1 | + | +++ | Zhou et al., 2015 | |
| BMP4 | +++ | ++ | + | Rege et al., 2015 |
| SIGNALING | ||||
| *†LEF1 | +++ | +/− | Rege et al., 2014 | |
| *†NOV | +++ | +/− | ||
| †SLC27A2 | + | +++ | ||
| †TSPAN12 | + | +++ | ||
| NGFIB | +++ | ++ | Bassett et al., 2004b | |
| Nurr-1 | +++ | + | ||
| PPARγ | ++ | + | Uruno et al., 2011 | |
| Ucn3 | + | + | Fukuda et al., 2005 | |
| CRF1 | +/− | ++ | ++ | |
| CRF2 | +/− | ++ | ++ | |
| LHR | +/− | + | Pabon et al., 1996 | |
| DETOXIFICATION | ||||
| Seladin | − | +++ | +++ | Battista et al., 2009 |
| †AKR1C3 | + | +++ | Rege et al., 2014 | |
This table lists some of the genes and gene products that have been used to define the different functions of the zones of the adrenal cortex, mostly using either immunocytochemistry (shown in normal text) or in situ hybridization (in italics). It is by no means exhaustive, many more components have been listed, notably when cDNA arrays have been used, (e.g., de Mendonca et al., 2014; Nishimoto et al., 2014; Tyczewska et al., 2014). Plus signs (+) indicate presence, minus signs (−) absence. The number of + signs gives a rough indication of the abundance of signal under control conditions, i.e., with no special treatment. Naturally all of these values may change, sometimes enormously, if neutral physiological regimes are perturbed.
*
Also reported in glomerulosa, but data not provided.
†
Transcriptome profiling.