(Halleraker et al., 1994). As sheep are only exposed to exogenous nonself antigens after birth because of the protection provided by a syndesmochorial placenta (Brambell, 1970;Boyd et al., 1976), it follows that factors other than exogenous nonself antigens are responsible for this modification.The interdependence of lymphoid-cell differentiation and stromal-cell development has been the subject of a number of studies. In man, rat and sheep, the early occurrence of FDCs was concomitant with B-cell aggregation and the appearance of primary follicles (Markgraf et al., 1982;Kroese et al., 1987;Halleraker et al., 1994 (Press et al., 1996) has allowed the investigation of the development of lymphoid and accessory-cell populations in a species shielded from exogenous antigen during gestation. In addition to FDCs in primary follicles, other accessory-cell populations that share a close association with B cells include the FDCs of gut-associated lymphoid follicles and macrophages of the marginal zone of the spleen. The aim of the present study was to characterize, using a panel of enzyme and immunohistochemical techniques, the accessory-cell populations in the lymphoid tissues of B-cell-depleted fetal sheep.
RESULTS
Ileal Peyer's Patch FolliclesThe small follicles in the ileal Peyer's patch of the Bcell-depleted fetuses demonstrated intense reactivity for the enzyme 5'nucleotidase (5'N), which was similar to the intensity of reactivity shown in the ileal Peyer's patch follicles of the untreated litter mates and the age-matched control fetuses (Figure 1). There was intense reactivity for magnesium-dependent adenosine triphosphatase (MgATPase) in some cells of the dome and interfollicular areas in the ileal Peyer's patch of the B-cell-depleted fetuses ( Figure 1). The dome region of ileal Peyer's patch of the Bcell-depleted fetuses showed increased reactivity for acid phosphatase whereas there was an absence of reactivity in the rudimentary follicle (Figure 1) There was also some reactivity for nonspecific esterase in the dome and interfollicular areas (not shown).A double immunofluorescence technique showed that the rudimentary follicles of the ileal Peyer' s patch in the B-cell-depleted fetuses showed strong staining for complement receptor-2 (CD21) and did not stain for IgM (not shown).
Primary Follicles in the Spleen and Lymph
NodesThe spleen and lymph nodes of the B-cell-depleted fetuses did not show the characteristic patterns of enzyme reactivity for 5'N, MgATPase, nonspecific esterase, and alkaline phosphatase that are associated with primary follicles (Figure 1). Serial step sectioning of the spleen and lymph nodes combined with staining for 5'N reactivity did not identify primary follicles in the B-cell-depleted fetuses. Primary follicles were readily identifiable by their typical pattern of enzyme reactivities in the control fetuses (Figure 1).The spleen and lymph nodes of B-cell-depleted fetuses contained a small population of CD21 cells.In the lymph nodes, most of these cells were scattered in the cortex, b...