Ultramicroscopic Examination of the Ovine Tonsillar Epithelia

As solid morphological knowledge of ovine tonsillar epithelia might contribute to a better understanding of the pathogenesis of several diseases including prion diseases, the epithelia of all tonsils of 7 one‐year‐old Texel sheep were examined using scanning and transmission electron microscopy. Major parts of the pharyngeal and tubal tonsils were covered by pseudostratified columnar ciliated epithelia that were interrupted by patches of epithelium containing cells with densely packed microfolds or microvilli, and cells with both microvilli and cilia. Smaller parts were covered by either flattened polygonal cells with densely packed microvilli or microfolds, squamous epithelial cells, or patches of reticular epithelium. The palatine and paraepiglottic tonsils were mainly lined by squamous epithelial cells with apical microplicae or short knobs. Additionally, regions of reticular epithelium containing epithelial cells with apical microvilli were seen. The lingual tonsil was uniformly covered by a keratinized squamous epithelium and devoid of microvillous cells and patches of reticular epithelium. The rostral half of the tonsil of the soft palate was lined by a pseudostratified columnar ciliated epithelium with characteristics of the pharyngeal and tubal tonsils. The epithelium of the caudal part resembled the epithelia of the palatine and paraepiglottic tonsils. Putative M cells, mainly characterized by apical microvilli or microfolds and a close association with lymphoid cells, seem manifestly present on the nasopharyngeal tonsils. The reticular epithelium of the palatine and paraepiglottic tonsils also harbor cells with small apical microvilli. The exact nature of these presumptive M cells should, however, be elucidated in functional studies. Anat Rec, 2010. © 2010 Wiley‐Liss, Inc.     

Yak (Bos grunniens) Tonsils: Morphological Description and Expression of IgA and IgG

This study aimed to describe the morphology, expression of IgA and IgG in adult yak tonsils. The 12 clinically healthy yak tonsils [3- to 6-year old, n = 12] were examined for morphology using light, and transmission electron microscopes. Expression of IgA and IgG was measured by qRT-PCR, ELISA, and immunohistochemistry. The results showed that the palatine tonsil, the tonsil of the soft palate, and the lingual tonsil were oropharyngeal tonsils. The stratified squamous epithelia covering them had a thick underlying layer of connective tissue and their crypts were heavily infiltrated by lymphocytes. The pharyngeal tonsil and the tubal tonsil were nasopharyngeal tonsils. The epithelia of them was predominantly pseudostratified columnar ciliary epithelium, which were loosely arranged with a number of desmosomes or intermediate junctions variably connecting them. The expression levels of IgA and IgG mRNA and protein from high to low was in the pharyngeal tonsil, palatine tonsil, tonsil of the soft palate, lingual tonsil, and tubal tonsil, respectively. Interestingly, the expression of IgG was very significantly higher than that of IgA in yak tonsils (P < 0.01). Both the IgA and IgG ASCs were distributed in the subepithelial areas of the non-reticular crypt epithelium, especially areas of pseudostratified columnar ciliary epithelium, the reticular crypt epithelium, lymphoid follicles, interfollicular areas, and with some of the positive cells aggregating around the glands. The results indicated that the tonsils were not only typical secondary lymphoid organs but also lymphoepithelial structures. IgG could be a significant component of mucosal immune responses in yak tonsils. Anat Rec, 302:999–1009, 2019. © 2018 Wiley Periodicals, Inc.     

Locations of gut-associated lymphoid tissue in the 3-month-old chicken: a review

The lymphoid tissue that is associated with the intestinal tract, the so-called gut-associated lymphoid tissue (GALT), is well developed in the chicken. Depending on the location, it is present as aggregations of lymphoid cells, or organized in lymphoid follicles and tonsils. From proximal to distal, the intestinal tract contains a pharyngeal tonsil, diffuse lymphoid tissue and lymphoid follicles in the cervical and thoracic parts of the oesophagus, an oesophageal tonsil, diffuse lymphoid tissue in the proventriculus, a pyloric tonsil, Peyer's patches, Meckel's diverticulum, two caecal tonsils, diffuse lymphoid tissue in the rectum, the bursa of Fabricius, and diffuse lymphoid tissue in the wall of the proctodeum. The lymphoid tissues are frequently covered by a lympho-epithelium that is infiltrated by lymphoid cells. Such an epithelium often contains M or microfold cells, which are specialized in antigen sampling and transport antigens to the underlying lymphoid tissue. A solid knowledge of the avian GALT could contribute to the development of vaccines to be administered orally. Additionally, immune stimulation via pre- and probiotics is based on the presence of a well-developed intestinal immune system.     

The potential role of the ovine pharyngeal tonsil in respiratory tract immunity: a scanning and transmission electron microscopy study of its epithelium

The mucosal epithelium of the pharyngeal tonsil was studied in 10 conventionally raised sheep by scanning and transmission electron microscopy. The surface of the tonsil was covered by two types of topographically and ultrastructurally distinct epithelium. The epithelium overlying the lymphoid follicle region (follicle-associated epithelium) was compressed by adjacent ciliated epithelium and consisted of predominantly non-ciliated cells of varying height. The majority of these cells possessed microvilli or microfolds of varying number, height and density on their surface. Small numbers of squamous cells and, occasionally, intermediate cells were also present in these areas. Ultrastructurally, the microvillous cells showed features similar to M cells which occur in other mucosal sites. They contained cytoplasmic vesicles and vacuoles and formed intercellular digital junctions. Between these cells was a heavy infiltration of lymphoid cells. Focal disintegration of the mucosal epithelium was occasionally seen. The epithelium covering non-follicle areas consisted of mainly ciliated cells interspersed with some goblet cells and squamous cells, morphologically resembling those elsewhere in the respiratory tract. These findings have provided evidence which confirms that the ovine pharyngeal tonsil is part of the respiratory tract-associated lymphoid tissue.     

Respiratory tract-associated lymphoid tissue in conventionally raised sheep

The distribution and morphological features of respiratory tract-associated lymphoid tissue was investigated in 20 clinically healthy sheep aged from 6 months to 9 years. Five different forms of lymphoid tissue were identified, but their distribution varied between regions of the respiratory tract and between individuals. Scattered lymphoid cells were the most predominant form. Dense lymphoid aggregations were frequently seen in the pharyngeal tonsil and bronchioles. They were less common in the mid-nasopharynx and about the opening of the auditory tube, occasionally seen in the anterior nasopharynx, trachea and bronchus and rarely found in the regions of the nasal conchae, nostril, nasal vestibule and epiglottis. Nodular lymphoid aggregations with morphologically distinct lympho-epithelium were seen only in the pharyngeal tonsil and opening of the auditory tube region. Small numbers of intra-epithelial lymphoid cells were always present throughout the mucosa, but intraluminal lymphocytes were observed mainly in the regions of pharyngeal tonsil, trachea, bronchi and large bronchioles. The respiratory tract-associated lymphoid tissue is less developed in sheep than in most other species studied, except perhaps cattle.     

Intercellular and lymphatic pathways associated with tonsils of the soft palate in young pigs

 Tonsils of the soft palate of pigs are the main oropharyngeal lymphoid tissues that protect the body against antigens entering through the mouth. The aim of this work was to elucidate the intercellular and lymphatic pathways by which lymph and cells are transported through these tonsils. Tonsillar tissue from freshly-killed pigs was examined using light microscopy and electron microscopy, or was injected with Mercox for scanning electron microscopy of corrosion casts. Intercellular fluid passes between epithelial cells and is continuous with that of the subepithelium. Fluid from the subepithelium flows into sinuses that form a network around the apex of follicles. These sinuses are continuous with parafollicular sinuses that penetrate the parafollicular tissue between the follicles. Some parafollicular sinuses are traversed by a complex network of cell processes, whereas others appear to lack such processes. Some parafollicular sinuses are closely located (10 μm) to venules; others lie adjacent to the follicle capsule. No lymphatics enter or leave the follicles. All lymph from the tonsils must traverse parafollicular sinuses before entering septal vessels, and these are continuous with basal vessels. Basal vessels coalesce to form efferent vessels that transport lymph from the tonsil to the primary lymph nodes. Septal, basal and efferent lymphatic vessels contain prominent valves and many lymphocytes. Lymphatic sinuses appear to be a significant pathway for lymphocytes migrating from the tonsillar lymphoid tissue. Accepted: 5 November 1997     

Immunoglobulin-containing cells in human tonsils as demonstrated by immunohistochemistry.

Intracellular immunoglobulin has been demonstrated in human palatine tonsils by the unlabelled antibody peroxidase-antiperoxidase complex (PAP) method in which rabbit antiserum to a range of human immunoglobulins (Igs) was linked to the PAP complex by an intermediate stage of swine antiserum to rabbit Ig. The effects of different methods of fixation and processing have been compared, formol-saline fixation giving the best results. The PAP technique proved greatly superior to the fluorescein isothiocyanate (FITC)-based technique, not only in sensitivity but in permitting study of the finer histological and cytological features. The lymphoid follicles are shown to have three distinct zones, two forming the follicle centre (zones (a) and (b)), and the third (zone (c)) the lymphocyte cap. Ig synthesis appeared to begin in the cells in zone (b). IgG, IgA, IgM, IgE and IgD were present in all tonsils, with IgG predominating, confirming that the tonsil resembles lymph nodes more closely than it does alimentary lymphoid tissue. Some follicles contained more than one type of Ig. The tonsil appears to have a well-developed T-dependent area, the lymphoid follicles forming a B-cell area. The structure of the tonsil would seem to facilitate contact between its lymphoid tissue and antigens in the crypts, and it is postulated that some T cells within the crypt epithelium, after contact with antigen, may leave the tonsil by the efferent lymphatics and enter the peripheral circulation by the thoracic duct, whilst other primed T cells interact with B cells in the follicle centres. Some B cells may then start to synthesize immunoglobulin, whilst others become memory cells in the lymphocyte 'cap' of the follicle.     

Development and immunophenotyping of the pharyngeal tonsil (adenoid) in cattle.

Immunoperoxidase staining and electron and light microscopy were used to characterize the development of the pharyngeal tonsil in 98 cattle aged between 30 days of gestation and 12 years. The rugae of the pharyngeal tonsil were poorly formed before 95 days of gestation. Microvillous (M) cells associated with intra-epithelial leucocytes (lympho-epithelium) were scattered among ciliated and goblet cells covering most of the surface in post-natal animals. Intra-epithelial leucocytes were rare in fetuses, but ciliated and M cells could be distinguished. Leucocytes of the lamina propria started to accumulate at approximately 120 days of gestation. A loose accumulation of mononuclear cells progressed into a B-cell rich upper and T-cell rich lower layer, with typical lymphoid tissue organization in post-natal animals and lymphoid involution in aged cattle. Primary lymphoid follicles formed at 5 months of gestation, but germinal centres did not form until 2 to 4 weeks after birth. Except for null cells, the relative number of cells staining for each leucocyte phenotype or MHC class II antigen increased with age, especially during the neonatal period. The early development, strategic location and specialized structure of the pharyngeal tonsil suggest an important role in modulating inhaled antigens in cattle. Fetal and neonatal calves had minimal lymphoid tissue priming, as indicated by lack of secondary follicles, low MHC class II expression and few intra-epithelial leucocytes. The phenotypic differences may be relevant to the increased susceptibility of calves to infectious diseases shortly after birth.     

Human tonsil intraepithelial B cells: a marginal zone-related subpopulation.

AIMS: To determine if intraepithelial B cells in reactive human palatine tonsils were similar to the marginal zone cells of the spleen and Peyer's patches. METHODS: Reactive human palatine tonsils were studied using conventional methods of light microscopy, electron microscopy, and a panel of monoclonal antibodies for leucocyte common antigens. RESULTS: Clinically important numbers of marginal zone-related B cells around the mantle zone were absent in lymphoid follicles, but in the cryptal epithelium there were abundant lymphoid cells with centrocyte-like nuclei and clear cytoplasm, intermingled with macrophages and plasma cells. The immunophenotype of these intraepithelial B cells was distinctive and similar to that found in the splenic marginal zone cells (IgM+, IgD-, CD23-, CD10-, CD35+, CD21+, bc12+, KB61+). CONCLUSIONS: Intraepithelial B cells in human tonsil could represent the counterpart of the marginal zone described in Peyer's patches. Their presence within the epithelium could reflect the destination for the malignant B cells in the lymphoepithelial lesion of mucosa associated lymphoid tissue (MALT) lymphomas. Human palatine tonsil lymphoid tissue has morphological, immunophenotypic, and pathological features similar to those of MALT.     

Immunohistochemical detection of prion protein in lymphoid tissues of sheep with natural scrapie.

The scrapie-associated form of the prion protein (PrPSc) accumulates in the brain and lymphoid tissues of sheep with scrapie. In order to assess whether detecting PrPSc in lymphoid tissue could be used as a diagnostic test for scrapie, we studied the localization and distribution of PrPSc in various lymphoid tissues collected at necropsy from 55 sheep with clinical scrapie. Samples collected from the spleen, palatine tonsil, ileum, and five different lymph nodes were immunohistochemically stained for PrPSc. PrPSc was found to be deposited in a reticular pattern in the center of both primary and secondary lymphoid follicles. In addition, granules of PrPSc were seen in the cytoplasm in macrophages associated with the lymphoid follicles. In 54 (98%) of the 55 scrapie-affected sheep, PrPSc was detected in the spleen, retropharyngeal lymph node, mesenteric lymph node, and the palatine tonsil. However, only in the palatine tonsils was PrPSc present in a consistently high percentage of the lymphoid follicles. PrP was not detected in any of the lymphoid tissues of 12 sheep that had no neurohistopathological signs of a scrapie infection. We conclude that the tonsils are the best-suited lymphoid tissue to be biopsied for the detection of PrPSc in the diagnosis of clinical scrapie in living sheep.     

Histological and immunohistological investigation of lymphoid tissue in the platypus (Ornithorhynchus anatinus)

The gross and histological appearance and the distribution of T and B lymphocytes and plasma cells are described for lymphoid tissues obtained from 15 platypuses. The spleen was bilobed and surrounded by a thick capsule of collagen, elastic fibres and little smooth muscle. White pulp was prominent and included germinal centres and periarterial lymphoid sheaths. Red pulp contained haematopoietic tissue. A thin lobulated thymus was located within the mediastinum overlying the heart. The cortex of lobules consisted of dense aggregates of small and medium lymphocytes, scattered macrophages and few reticular epithelial cells. In the medulla, Hassall's corpuscles were numerous, lymphocytes were small and less abundant, and reticular cells were more abundant than in the cortex. Lymphoid nodules scattered throughout loose connective tissue in cervical, pharyngeal, thoracic, mesenteric and pelvic sites measured 790±370 μm (mean±S.D. , n = 39) in diameter, the larger of which could be observed macroscopically. These consisted of single primary or secondary follicles supported by a framework of reticular fibres. Macrophages were common in the germinal centres. The platypus had a full range of gut-associated lymphoid tissue. No tonsils were observed macroscopically but histologically they consisted of submucosal follicles and intraepithelial lymphocytes. Peyer's patches were not observed macroscopically but histologically they consisted of several prominent submucosal secondary follicles in the antimesenteric wall of the intestine. Caecal lymphoid tissue consisted of numerous secondary follicles in the submucosa and densely packed lymphocytes in the lamina propria. Bronchus-associated lymphoid tissue was not observed macroscopically but was identified in 7 of 11 platypus lungs assessed histologically. Lymphoid cells were present as primary follicles associated with bronchi, as aggregates adjacent to blood vessels and as intraepithelial lymphocytes. The distribution of T lymphocytes, identified with antihuman CD3 and CD5, and B lymphocytes and plasma cells, identified with antihuman CD79a and CD79b and antiplatypus immunoglobulin, within lymphoid tissues in the platypus was similar to that described in therian mammals except for an apparent relative paucity of B lymphocytes. This study establishes that the platypus has a well-developed lymphoid system which is comparable in histological structure to that in therian mammals. It also confirms the distinctiveness of its peripheral lymphoid tissue, namely lymphoid nodules. Platypus lymphoid tissue has all the essential cell types, namely T and B lymphocytes and plasma cells, to mount an effective immune response against foreign antigens.     

Evidence of M cells as portals of entry for antigens in the nasopharyngeal lymphoid tissue of humans

The nasopharyngeal tonsils (adenoids) are prominent components of human nasal-associated lymphoid tissues (NALT). However, the role of the nasopharyngeal tonsils in antigen uptake for initiation of the mucosal immune response is unknown. The aims of this study were to describe the ultrastructure and function of the M cells of the human nasopharyngeal tonsils and to clarify their capacity for antigen uptake. Tissues obtained from eight patients undergoing adenectomy were examined by light and electron microscopy. Lymphoepithelium covers the nasopharyngeal lymphoid tissue and consists of ciliary epithelium, non-ciliary epithelial cells, M cells, goblet cells, and many intraepithelial lymphoid cells. M cells have irregular and broad cytoplasm-containing microvilli on their surface and small vesicles in their cytoplasm. Many lymphoid cells were enfolded by M cells. The uptake of horseradish peroxidase (HRP) in the tissue in organ culture was studied using histochemical techniques. Excised adenoid tissue was incubated in RPMI 1640 culture media with HRP for 10, 30, and 60 min. HRP which had adhered to the surface was taken up in vesicles and then transported in vesicles and tubules by M cells. The M cells of nasopharyngeal lymphoid tissue were ultrastructurally and functionally similar to those in human Peyer’s patches and colonic lymphoid follicles. These findings indicate that NALT bears similarities to the gut-associated lymphoid tissue, and its antigen uptake capacity may be important for initiation of immunity in the upper aerodigestive tract. Received: 8 July 1999 / Accepted: 17 December 1999     

Development of the palatine tonsil in conventional and germ-free piglets

Palatine tonsils, like the Peyer's patches, are considered to be major inductive sites for the mucosa-associated lymphoid tissue (MALT), providing sampling and effector functions for the upper respiratory tract. Consistent with this, they have the architecture required of a classic inductive site (B-cell follicles, immunoglobulin class switching and the presence of na?ve and memory T-cells). Here we show that much of this architecture develops after birth in the neonatal piglet, the numbers of T-cells, B-cells and accessory cells increasing with age. Conventional piglets also had higher levels of activated and memory T-cell subsets than germ-free piglets, consistent with development occurring as a result of microbial stimulus. The results suggest that the microbial environment influences the development of the tonsil immunological architecture. Given the role of the tonsil in induction of mucosal responses, this raises questions as to the effectiveness of the tonsil in dealing with colonising organisms in the neonate.     

M cells at locations outside the gut.

Lymphoid tissue associated with mucosal membranes is found not only along the gastrointestinal tract, but also in the tonsils, the upper and lower airways, and the conjunctiva of the eye. The epithelia overlying this mucosa-associated lymphoid tissue (MALT) contain membranous (M) cells which transport antigenic matter across the mucosal membrane to initiate immune responses. Although the morphology and function of intestinal M cells have been thoroughly studied, relatively little is known about the presence and properties of M cells in MALT outside the gut. The available data on ultrastructure, histochemistry, and antigen sampling function of the epithelia in tonsils, nasal-, larynx-, bronchus-, and conjunctiva-associated lymphoid tissue are reviewed and critically discussed. It is concluded that, in principle, the concepts of mucosal immune protection can be applied to these sites of MALT. However, it is questionable whether a separate cell type similar to intestinal M cells exists and performs antigen sampling in the different MALT epithelia. Further studies combining functional and morphological techniques are essential to understand the initiation of immune reaction at the mucosal membranes.     

An unusual structure of venules in tonsils of the soft palate of young pigs

In tonsils of the soft palate of pigs high endothelial venules (HEV) occur throughout the parafollicular region. In some regions of the venules the endothelial lining is uniformly high; in other areas, mainly the nuclear region of the endothelial cell protrudes into the lumen and the cytoplasmic region is attenuated. In this portion of the venule endothelial cells may traverse the lumen of the vessel to form intravascular bridging processes. The endothelial cells are characterised by pinocytotic vesicles, many surface microvilli, cytoplasmic processes and local cell-cell contacts with adjacent endothelial cells. These specialised HEV may play a role in the unusual pattern of lymphocytes from tonsillar tissue into blood. In the tonsils of the soft palate of pigs, HEV may mediate both lymphocyte migration to and from the lymphoid tissue. Further investigation is required to determine the importance of these 2 migration pathways.     

Immunohistochemical detection of co-localizing cytokine and antibody producing cells in the extrafollicular area of human palatine tonsils.

In vitro experiments have documented the role of cytokines in the regulation of the human humoral immune response. Which cytokines are operative in vivo and in which lymphoid compartment interactions between cytokine-producing T cells and antibody-forming B cells occur is still unclear. For that reason we studied human tonsils using immunohistochemical techniques. In tissue sections from tonsils in a resting stage after recurrent tonsillitis we observed cells producing IL-1 alpha and tumour necrosis factor-alpha (TNF-alpha) which were exclusively localized in the mantle zone of the follicle and in the extrafollicular area. Furthermore, a high frequency of interferon-gamma (IFN-gamma)-producing cells was detected in the extrafollicular area, but not inside the follicles. Occasional IL-2- and IL-4-producing cells were found in the extrafollicular area. Immunohistochemical detection of antibody isotypes revealed that B cells, IgM-membrane-positive, were localized inside the follicles and mantle zones, whereas IgD-membrane-positive cells were mainly found in the mantle zones of secondary follicles. In contrast, plasma cells producing IgG1-4 and IgA1-2 were found in the extrafollicular area. No IgD and IgE antibody-forming cells were detected in tonsils, whereas IgM antibody-forming cells were detected in the extrafollicular area. The co-localization of cytokine-producing cells and antibody-forming cells in human tonsil suggests that T-B cell interactions, required for B cell differentiation and isotype switching, take place in the extrafollicular area.     

The association of sore throat and psoriasis might be explained by histologically distinctive tonsils and increased expression of skin‐homing molecules by tonsil T cells

Recent studies have highlighted the involvement of the palatine tonsils in the pathogenesis of psoriasis, particularly among patients with recurrent throat infections. However, the underlying immunological mechanism is not well understood. In this study we confirm that psoriasis tonsils are infected more frequently by β‐haemolytic Streptococci, in particular Group C Streptococcus, compared with recurrently infected tonsils from patients without skin disease. Moreover, we show that tonsils from psoriasis patients contained smaller lymphoid follicles that occupied a smaller tissue area, had a lower germinal centre to marginal zone area ratio and contained fewer tingible body macrophages per unit area compared with recurrently infected tonsils from individuals without skin disease. Psoriasis patients' tonsils had a higher frequency of skin‐homing [cutaneous lymphocyte‐associated antigen (CLA⁺)] CD4⁺ and CD8⁺ T cells, and this correlated significantly with their frequency of blood CLA⁺ T cells. The psoriasis patients also had a higher frequency of tonsil T cells expressing the interleukin (IL)‐23 receptor that was expressed preferentially by the CLA⁺ T cell population. In contrast, recurrently infected tonsils of individuals without skin disease had a higher frequency of tonsil T cells expressing the activation marker CD69 and a number of chemokine receptors with unknown relevance to psoriasis. These findings suggest that immune responses in the palatine tonsils of psoriasis patients are dysregulated. The elevated expression of CLA and IL‐23 receptor by tonsil T cells may promote the egression of effector T cells from tonsils to the epidermis, suggesting that there may be functional changes within the tonsils, which promote triggering or exacerbation of psoriasis.     

Fine structure of the anal tonsilliar epithelium in the laboratory shrew (Suncus murinus)

The epithelium of the anal tonsil of the laboratory shrew was studied by scanning and transmission electron microscopy, with particular attention focused on the structure of the epithelium lining the anal tonsillar crypt. The tonsillar crypt surface is lined by two kinds of epithelia: squamous epithelium, which is located mainly at the neck of the crypt and includes keratohyalin granules in the superficial layer, and reticular epithelium, which is invaded by many immigrating cells and has several micropores immigrating cells to pass through. In addition, basal granulated cells are present in the basal layer. These results suggest that the reticular epithelium of the anal tonsil belongs to the well-developed gut-associated lymphoid tissue (GALT) in the alimentary canal. It represents a specialized and important compartment in immunological function, similarly to the palatine tonsils of other mammals, and has as yet unknown roles in digestion.     

The enzyme histochemistry of lymphoid and non-lymphoid cells of the human palatine tonsil: a basis for the study of lymphomas

The distribution of various hydrolytic enzymes has been determined in 27 human palatine tonsils by means of conventional enzyme histochemical techniques, and lysozyme (muramidase) activity has been localised in eight tonsils by the unlabelled antibody peroxidase-antiperoxidase complex (PAP) method. The enzyme activities of various cell types are compared and the effect of various methods of fixation and processing discussed. The results suggest that arrangement of various histiocytic cells types within the tonsillar follicles and crypt epithelium is related to the processing of antigen. The PAP method for lysozyme demonstrates a smaller population of cells than is demonstrated by the alpha-naphthyl acetate esterase (ANAE) method. T-cells are demonstrated by the presence of dot-like ANAE activity in their cytoplasm. Large numbers of the lymphocytes of this type were located in the T-dependent areas of the tonsil, and are frequent beneath the crypt epithelium. The efferent lymphatic vessels appeared to contain an almost pure population of T-cells. The immunohistochemical method for lysozyme did not differentiate between T- and B-cell areas, dot-like activity being absent. As in other workers' studies on non-lymphoid cells of the murine spleen, several types of glass-adherent cells have been identified in short-term cell cultures from the human tonsil. True dendritic cells and branching macrophages differ in several ways (as in the mouse spleen). The tonsil is considered to be a useful control "reactive" lymphoid organ, to act as a baseline tissue in an extended study of morphology and enzyme histochemistry in the lymphomas.