Milán FJ and Puelles L (2000), Patterns of calretinin, calbindin, and tyrosine...

XB-ART-11347

J Comp Neurol 2000 Mar 27;4191:96-121.

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Patterns of calretinin, calbindin, and tyrosine-hydroxylase expression are consistent with the prosomeric map of the frog diencephalon.

Milán FJ , Puelles L .

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This paper re-examines a previously published segmental map of the frog diencephalon (Puelles et al. [1996] Brain Behav.Evol. 47:279-310) by means of immunocytochemical mapping of calretinin, calbindin, and tyrosine hydroxylase. The distribution of neuronal populations, axon tracts, and neuropils immunoreactive for these markers was studied in adult specimens of Rana perezi and Xenopus laevis sectioned sagittally or horizontally. Emphasis was placed on study of the relationship of observed chemoarchitectural boundaries with the postulated overall prosomeric organization and the schema of nuclear subdivisions we reported previously, based on acetylcholinesterase histochemistry and Nissl pattern in Rana. The data reveal a large-scale correspondence with the segmental map in both species, although some differences were noted between Rana and Xenopus. Notably, retinorecipient neuropils were generally immunoreactive for calretinin only in Rana. Importantly, calretinin immunostaining underlines particularly well the transverse prosomeric boundaries of the dorsal thalamus. A number of nuclear subdivisions noted before with AChE were corroborated, and some novel subdivisions became apparent, particularly in the anterior nucleus of the dorsal thalamus and in the habenular complex. The mapping of tyrosine hydroxylase clarified the segmental distribution of the catecholaminergic cell groups in the frog forebrain, which is comparable to that observed in other vertebrates.

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Species referenced: Xenopus laevis
Genes referenced: ache calb1 calb2 tbx2 tff3.7

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	Fig. 1. Lateral sagittal sections of a Xenopus brain immunoreacted for calretinin, shown in lateromedial order (A–C). Dorsal is oriented upward and rostral to the left, as in all other sagittal sections shown in this paper. The diencephalic prosomeres p1–p3 and the isthmus are delimited by dash lines. The alar-basal boundary in p2 and p3 passes just under the horizontal course of the thalamotelencephalic tract (tht) as it extends from the dorsal thalamus to the root of the lateral forebrain bundle in the secondary prosencephalon (best seen in A and B). Scale bar 5 1 mm in C (also applies to A,B).
	Fig. 2. Adjacent lateral sagittal sections of the same brain as in Figure1, but immunoreacted for calbindin. Each lateromedial level (A–C) corresponds to one of the CR-stained ones in Figure 1. Scale bar 5 1 mm in C (also applies to A,B).
	Fig. 3. A,B: Continuation of the CR-immunoreacted sagittal section series begun in Figure 1, to be compared with Figure 4A,B. A is lateral to B. Transverse diencephalic interprosomeric boundaries and the isthmus are indicated by dashed lines as in Figure 1. C: Detail at higher magnification of CB in the habenular region in a section slightly medial to that in B. Note the CB-IR neurons in the dorsocaudal thalamic area (DC), which separate the habenular complex from the pretectum (p1). Scale bar 5 250 mm in C (1 mm for A,B).
	Fig. 4. A,B: Sagittal sections adjacent to those in Figure 3A,B, immunoreacted for calbindin. C: Higher magnification detail of CB in the mammillary and retromammillary regions in a sagittal section slightly medial to B. D: Higher magnification detail of CB in the hypothalamus in a section medial to C: note CB-IR fibers possibly originating in the preoptic magnocellular nucleus (PMg) passing through the retrochiasmatic periventricular area into the tuberal hypothalamus; the median floor area of the mammillary, retromammillary, and tuberculum posterior regions shows intense cellular and neuropil staining. Scale bar 5 250 mm in D (also applies to C; 1 mm for A,B).
	Fig. 7. Horizontal sections of Xenopus or Rana brains immunoreacted for calretinin. Rostral is oriented to the bottom of the page. A: Xenopus. Dorsal section passing through the habenular region and the caudal poles of the telencephalon. B: Rana/Xenopus (corresponding halves are shown for comparison; identified at the top). Sections at dorsal levels of the neuropil of Bellonci (compare with schema in Fig. 5B). Note staining differences in the retinorecipient neuropils. Curiously, the lateral thalamic neuropil (LN) is more CR-IR in Xenopus than in Rana. Scale bar 5 250 m in B (also applies to A).
	Fig. 8. A: Rana/Xenopus (corresponding halves are shown for comparison; identified at the top; orientation as in Fig. 7). Horizontal CR-immunoreacted sections at level schematized in Figure 5C, passing through the lower part of the neuropil of Bellonci. Note differences in retinorecipient neuropils. B: Xenopus. Horizontal CRimmunoreacted section ventral to the neuropil of Bellonci. Scale bar 5 250 mm in B (also applies to A).
	Fig. 9. A: Xenopus. Detail of horizontal CR-immunoreacted section through the dorsal thalamus and pretectum (PT), showing the bisbenzimide fluorescent counterstain used here to identify roughly the proportion of CR-IR cells in the dorsal thalamus. The ventricle lies at the right and the optic tract at the left. Note CR-negative cells building up mainly in the periventricular stratum (in contrast to the largely CR-negative PT). B, C: Xenopus/Rana (roughly corresponding halves shown for comparison; identified at the top). Horizontal CRimmunoreacted sections approximately at the ventralmost part of the diencephalic alar plate [note the fiber bundles of the thalamotelencephalic tract (tht) and the lateral forebrain bundle (lfb)]. Note again differences in retinorecipient neuropils, as well as in the suprachiasmatic and preoptic regions. Scale bar 5 250 mm in C (also applies to A,B).
	Fig. 10. A–D: Continuation of the dorsoventral series of horizontal CR-immunoreacted sections through the Xenopus brain illustrated in Figures 7–9. A: Section at the level of the optic chiasma (oc), showing the transition from the diencephalic alar plate into the corresponding basal plate, which has a quite different expression pattern (note horizontal course of the tht tract). B–D: Successively more ventral section levels through the hypothalamus, ending at the mammillary nucleus. E: Sagittal CR-immunoreacted section through a Rana brain; rostral to the left. The plane passes just through the thin thalamoeminential tract (compare Figs. 7B and 8A), illustrating the slight dorsoventral dispersion of its component fiber bundles. Note as well the CR-IR uncinate neuropil and basal optic nucleus (U; BON). Scale bar 5 250 mm in A–D; 1 mm in E.
	Fig. 11. A: Higher magnification detail of a sagittal section lying in between Figure 1A,B, showing the CR-IR thin fibers of the thalamohypothalamic tract (thh) as they course out of the p2 alar plate (dorsal thalamus) and enter the underlying basal plate. Note the contrasting change of course of the thalamotelencephalic fibers (tht), which are more immunoreactive. B: Higher magnification detail of CR-IR cells and fibers of the magnocellular nucleus of the posterior commissure, as observed in a sagittal section, also containing the adjoining superficial griseum tectale formation of the midbrain (GTs). C: Higher magnification detail of the CR-IR core of the neuropil of Bellonci in a sagittal section in Xenopus. Rostral is oriented to the left. Scale bar 5 300 mm in C (also applies to A,B).